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Zoological Journal of the Linnean Society, 2020, XX, 1–33. With 10 figures. Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020

A revision of pipistrelle-like bats (Mammalia: Chiroptera: Vespertilionidae) in East Africa with the description of new genera and species

ARA MONADJEM1,2,3, , TERRENCE C. DEMOS3, DESIRE L. DALTON1,4, PAUL W. WEBALA5, SIMON MUSILA6, JULIAN C. KERBIS PETERHANS3,7 and BRUCE D. PATTERSON3

1Department of Biological Sciences, University of Eswatini, Private Bag 4, Kwaluseni, Eswatini 2Mammal Research Institute, Department of Zoology and Entomology, University of Pretoria, Private Bag 20, Hatfield 0028, Pretoria, South Africa 3Negaunee Integrative Research Center, Field Museum of Natural History, Chicago, IL 60605, USA 4South African National Biodiversity Institute, PO Box 754, Pretoria 0001, South Africa 5Department of Forestry and Wildlife Management, Maasai Mara University, Narok 20500, Kenya 6Mammalogy Section, Zoology Department, National Museums of Kenya, PO Box 40658-00100, Nairobi, Kenya 7College of Arts & Sciences, Roosevelt University, 430 S Michigan, Chicago, IL 60605, USA

Received 29 May 2020; revised 1 July 2020; accepted for publication 4 July 2020

Vespertilionidae (class Mammalia) constitutes the largest family of bats, with ~500 described species. Nonetheless, the systematic relationships within this family are poorly known, especially among the pipistrelle-like bats of the tribes Vespertilionini and Pipistrellini. Perhaps as a result of their drab pelage and lack of obvious morphological characters, the genus and species limits of pipistrelle-like bats remain poorly resolved, particularly in Africa, where more than one-fifth of all vesper bat species occur. Further exacerbating the problem is the accelerating description of new species within these groups. In this study, we attempt to resolve the systematic relationships among the pipistrelle-like bats of sub-Saharan Africa and Madagascar and provide a more stable framework for future systematic efforts. Our systematic inferences are based on extensive genetic and morphological sampling of > 400 individuals covering all named genera and the majority of described African pipistrelle-like bat species, focusing on previously unstudied samples of East African bats. Our study corroborates previous work by identifying three African genera in Pipistrellini (Pipistrellus, Scotoecus and Vansonia), none of which is endemic to Africa. However, the situation is more complex in Vespertilionini. With broad taxonomic sampling, we confirm that the genus Neoromicia is paraphyletic, a situation that we resolve by assigning the species of Neoromicia to four genera. Neoromicia is here restricted to Neoromicia zuluensis and allied taxa. Some erstwhile Neoromicia species are transferred into an expanded Laephotis, which now includes both long- eared and short-eared forms. We also erect two new genera, one comprising a group of mostly forest-associated species (many of which have white wings) and the other for the genetically and morphologically unique banana bat. All four of these genera, as recognized here, are genetically distinct, have distinctive bacular morphologies and can be grouped by cranial morphometrics. We also demonstrate that the genus Nycticeinops, until now considered monospecific, includes both Afropipistrellus and the recently named Parahypsugo, thus representing the fifth African genus in Vespertilionini. A sixth genus, Hypsugo, is mostly extra-limital to sub-Saharan Africa. Finally, we describe three new species of pipistrelle-like bats from Kenya and Uganda, uncovered during the course of systematic bat surveys in the region. Such surveys are greatly needed across tropical Africa to uncover further bat diversity.

ADDITIONAL KEYWORDS: Africa – alpha taxonomy – genus revision – Mammalia – mitochondrial DNA – new genera – new species.

*Corresponding author. E-mail: [email protected] [Version of record, published online 10 September 2020; http:// zoobank.org/ urn:lsid:zoobank.org:pub: 71737F08-2938-4403- 8385-5438B2E5EABE] © 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 1 2 A. MONADJEM ET AL.

INTRODUCTION attention (Monadjem et al., 2020b). Previous studies have suggested that it might be paraphyletic (Koubínová Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 Vespertilionidae is the largest chiropteran family et al., 2013; Monadjem et al., 2013; Goodman et al., in the world, with ~500 described species in 54 2015), and a close relationship between the distinctly genera (Moratelli & Burgin, 2019), of which 17 long-eared Laephotis Thomas, 1901 species and some genera and ≥ 106 species have been reported from members of the genus Neoromicia has been reported Africa (AfricanBats NPC, 2019; Monadjem et al., (Hoofer & Van den Bussche, 2003; Görföl & Csorba, 2020b). Within this group, the taxonomy of the 2018). Furthermore, the placement of Neoromicia nana pipistrelle-like or ‘pipistrelloid’ tribes Pipistrellini (Peters, 1852) (sometimes previously called Pipistrellus and Vespertilionini (sensu Amador et al., 2018), nanus or Hypsugo nanus) has been problematic. For subfamily Vespertilioninae, has been particularly example, unlike other species placed in Neoromicia, difficult to resolve. The systematic relationships of Neo. nana has a distinctive character set: the first these bats have been the subject of much debate over upper premolar is present, the braincase is obviously the past few decades (Hill & Harrison, 1987; Volleth and highly inflated, there is a distinct thumbpad & Heller, 1994; Hoofer & Van den Bussche, 2003; present at the base of the thumb, and the lower third Roehrs, Lack & Van den Bussche, 2010; Koubínová molar is ‘nyctalodont’ (Monadjem et al., 2010; Van et al., 2013; Amador et al., 2018; Moratelli & Burgin, Cakenberghe & Happold, 2013). Another taxonomic 2019). A well-supported phylogeny, particularly problem relates to the newly described Afrotropical of basal nodes, has not yet been published, and genus Parahypsugo, which is rendered paraphyletic by although species limits are poorly known for many Pipistrellus (Afropipistrellus) grandidieri Dobson, 1876 taxa (Andriollo et al., 2015), new species and genera (Monadjem et al., 2020a), and the relationship of these continue to be described (Benda et al., 2016; Hutterer two groups has not yet been investigated with respect to et al., 2019; Görföl et al., 2020). Nycticeinops Hill & Harrison, 1987. The high diversity of pipistrelle-like bats in Africa In contrast, the systematic relationships in has been overshadowed by the ambiguity of species Pipistrellini are somewhat more clear (Hoofer & relationships and further complicated by the local or Van den Bussche, 2003; Amador et al., 2018). Sub- regional focus of previous analyses (Monadjem et al., Saharan Africa was previously thought to be home to 2013; Goodman et al., 2015, 2017; Hutterer et al., 2019). two genera, Pipistrellus and Scotoecus Thomas, 1901. For example, in the past decade, two new species have However, a recent study demonstrated that Pipistrellus been described in the genus Neoromicia Roberts, 1926 rueppellii J. Fischer, 1829 is sister to Pipistrellus (Monadjem et al., 2013; Decher et al., 2015), a third + Nyctalus + Glischropus (Koubínová et al., 2013), in Pipistrellus Kaup, 1829 (Monadjem et al., 2020a) rendering that concept of Pipistrellus paraphyletic. and a fourth in Parahypsugo Hutterer et al., 2019, all Based on this evidence and the distinctive morphology from the Upper Guinea forest zone of West Africa. The of Pip. rueppellii, this species has since been placed phylogenetic relationships of pipistrelle-like bats in this in its own genus, Vansonia (Roberts, 1946), which region have been investigated with mitochondrial DNA was formerly recognized as a subgenus of Pipistrellus markers, and a new genus (Parahypsugo) was recognized (Moratelli & Burgin, 2019). New species continue to be based on genetic and morphological characters (Hutterer described in this tribe (Monadjem et al., 2020a). & Kerbis Peterhans, 2019; Monadjem et al., 2020a). The main objectives of this study are as follows: (1) to Likewise, the species identities and relationships present a phylogeny for pipistrelle-like vespertilionids of pipistrelle-like bats in Madagascar have been (tribes Pipistrellini and Vespertilionini) in sub- re-examined (Goodman & Ranivo, 2004; Bates et al., Saharan Africa based on a unique dataset of > 400 2006; Goodman et al., 2012, 2015), as have the southern specimens that have been sequenced and examined African Neoromicia (Goodman et al., 2017). In contrast, morphologically; (2) to investigate the putative the pipistrelle-like bats of East Africa have received paraphyly within the genus Neoromicia and resolve almost no attention, and many basic taxonomic and this taxonomic problem; and (3) to use an integrative systematic questions remain unanswered (Patterson & taxonomic approach to describe three East African Webala, 2012). East Africa appears to be a hub of cryptic species new to science. diversity in other bat families and genera (Demos et al., 2018, 2019a, b, 2020; Patterson et al., 2018, 2019, 2020). Mizerovská et al. (2019) argued that this is true for non- volant mammal faunas, and we expect this to be true for MATERIAL AND METHODS pipistrelle-like bats also. Numerous systematic problems in pipistrelle-like Study sites bats remain unresolved, particularly in Vespertilionini. Most of the material newly reported in this study The genus Neoromicia has been the focus of much was collected in the course of systematic surveys in

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 3 eastern, central and southern Africa over the past Data collection Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 three decades by Field Museum scientists and a host Our field protocols involved extensive use of hand nets, of in-country collaborators. This work was centred harp traps, mist nets and triple-high net suspension on East Africa, which is remarkable for a number systems. Hand nets were used exclusively at day-roost of reasons. First, it is bisected by the Equator and, sites. We used both two-bank and four-bank Austbat consequently, is host to exceptional biodiversity in harp traps (Faunatech.com.au), in addition to a many taxonomic groups, especially bats (Patterson & discontinued model manufactured by Bat Conservation Webala, 2012). Bats comprise at least one-quarter of and Management (batmanagement.com), which also the megadiverse mammal faunas of Kenya and Uganda produced our three triple-high systems. On the ground, (Thorn & Kerbis Peterhans, 2009; Musila et al., 2019). we deployed 6 and 12 m nylon mist nets in likely Second, East Africa is a region of long-term tectonic flyways and monitored those continuously, typically activity that has created the tallest mountains and from dusk to about midnight. The position of all study deepest depressions in Africa (Spawls & Mathews, sites was marked using a Garmin eTrex Vista HCx 2012) and given rise to an unparalleled diversity of global positioning system, from which elevation was habitats. In addition to its endemic montane faunas, also read. Bats selected for further investigation were East Africa is where the great equatorial rain forest placed into individual cloth bags and transported to a of Africa reaches its eastern limits, the Sahel reaches portable flight cage (4 m × 4 m × 2 m in size) of cloth its south-eastern limits, the Horn of Africa reaches its draped over a jointed PVC frame. Once released, the south-western limits and the eastern savannas reach bats flew around the cage, searching for an exit, and their northern limits. The region lies at the nexus of their calls were recorded using a hand-held ultrasound several biodiverse biomes (Linder et al., 2012). Third, detector (Pettersson D1000X; Pettersson Elektronik the infrastructure and relative political stability of AB, Uppsala, Sweden; 384 or 500 kHz sampling rate, the region have permitted long-term scientific efforts. 16-bit resolution). For sound analysis, a customized Several museum scientists documenting the mammal 512-point fast Fourier transform (FFT) was used with faunas in this region have collected extensive and a Hanning window for both spectrograms and power largely complementary sets of specimens, including spectrum. Following Jung, Molinari & Kalko (2014), associated tissue samples for genomic work, for we characterized echolocation calls by measuring the understanding the regional diversity of the bat faunas peak frequency or frequency with maximum energy of Africa. The contributions of the late Bill Stanley (FME), maximum frequency (StartF) and minimum throughout Tanzania and Dr Robert Kityo and his frequency (EndF) using Kaleidoscope v.3.1.4b team of students from Makerere University in Uganda (Wildlife Acoustics, USA). The mean of ten calls with deserve special mention here (Kityo & Kerbis, 1996; the best signal-to-noise ratios was calculated for Stanley et al., 1996, 1998; Kerbis Peterhans et al., each bat. 1998; Stanley & Goodman, 2011). Procedures involving voucher specimens followed Kenya, which occupies a central position in the guidelines established in mammalogy (Sikes, 2016) region, has been the focus of extensive bat surveys and were approved by Field Museum’s Institutional since 2006, organized by the Field Museum of Natural Animal Care and Use Committee (2012-003). After History (FMNH) in Chicago in partnership with euthanasia with halothane, individual bats were the National Museums of Kenya (NMK), the Kenya fumigated in ethyl ether and carefully inspected for Wildlife Service, Karatina University and Maasai Mara ectoparasites, all of which were preserved as a lot University. The ‘Bats of Kenya’ project had as its goal in 95% ethanol for parallel studies by parasitologist the development of a comprehensive understanding of Carl W. Dick. The total length (head and body plus the bat diversity of Kenya, including the production tail), tail length, hindfoot length (including claw), ear of a vouchered reference call library. Once fieldwork pinna length (from notch) and tragus length (where began, it soon became apparent that existing keys (e.g. present) were measured with a ruler in millimetres. Patterson & Webala, 2012) were of limited value in Body mass was weighed using Pesola balances and cataloguing the diversity of East African bats, because recorded to the nearest 0.1 g. In most cases, the species were geographically too variable, real species pectoral muscle was exposed, and two 0.5 cm2 samples limits were unrecognizable, taxonomic names were of muscle were placed within 1–2 h after death into being misapplied, including in genomic databases, or a liquid nitrogen dewar. Bat carcasses labelled with all of the above acting in concert. We therefore focused individual field numbers were injected with a 10% efforts on thorough documentation of each species with formalin solution and immersed in that formalin which we came into contact. To do this, we travelled to solution for 4–20 days, before being rinsed and all accessible parts of the country, neglecting only its transferred to 70% ethanol. In most cases, the skulls border regions with South Sudan and Somalia. of alcohol-preserved bats were removed by skinning

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 4 A. MONADJEM ET AL. the head from the lips back to the neck, severing the Phylogenetic analysis Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 spinal column at the occipital condyle and removing We used jMODELTEST2 (Darriba et al., 2012) on the skull; by re-everting the skin of the head around CIPRES Science Gateway v.3.1 (Miller et al., 2010) a ball of cotton, the morphology of the ears, lips and to determine the sequence substitution models that nose was still apparent and could be studied. The best fit the Cytb data using the Bayesian information skull and mandible were then cleaned by dermestid criterion (BIC). Uncorrected Cytb sequence divergences beetles for study. Although this entire procedure (p-distances) among and within species were calculated was specific to the ‘Bats of Kenya’ project, most for Cytb using MEGA X v.10.1.7 (Kumar et al., 2018). parts of it other than the flight cage and bioacoustic Maximum likelihood (ML) analysis was performed with recordings were used in mammal surveys elsewhere. the software IQ-TREE v.1.6.10 (Nguyen et al., 2015; All specimens and associated tissue samples were Chernomor et al., 2016) on the CIPRES portal. Gene deposited in the mammal collections of FMNH, tree analyses using a Bayesian inference (BI) model NMK and other national repositories, according to were generated in MrBayes v.3.2.7 (Ronquist et al., agreements between collaborating institutions. 2012) on the CIPRES portal for the same alignment as the ML analysis. Two independent runs were conducted in MrBayes using four Markov chains run DNA extraction, amplification and sequencing for 1 × 108 generations under default heating values Genomic DNA from frozen tissue samples was and sampled every 1000th generation. A conservative extracted using the Wizard SV 96 Genomic DNA 20% burn-in was used, and stationarity of the results Purification System (Promega Corporation, WI, were assessed using TRACER v1.7 (Rambaut et al., USA) or the DNeasy Blood and Tissue Kit (Qiagen). 2018). Majority-rule consensus trees were assembled Specimens were sequenced for mitochondrial for each Bayesian analysis. cytochrome b 3(Cytb), using the primer pair LGL 765F and LGL 766R (Bickham et al., 1995, 2004). We generated original genetic data from 310 Craniodental morphology individuals collected at 110 georeferenced localities Eight cranial and four dental measurements were and complemented them with 108 mitochondrial taken with callipers to the closest 0.01 mm, following sequences from 63 unique localities downloaded Monadjem et al. (2013). The cranial measurements from GenBank; of these GenBank sequences, 28 of were as follows: greatest skull length (GSKL), from the the specimens associated with them were examined posteriormost point of the occipital to the anteriormost by AM. Sequence data were obtained for a total point of the incisors; condylo-incisive length (CIL), from of 58 species/putative species in Vespertilionidae. the occipital condyles to the anteriormost point of the All individuals were sequenced for Cytb in order canines; greatest zygomatic breadth (ZYGO), taken to maximize the assessment of genetic diversity. as the greatest width across the zygomatic arches; However, redundant haplotypes were removed for greatest braincase width (GBW), braincase width subsequent phylogenetic analyses (for complete list of taken in the frontal plane above the zygomatic arches; individuals sequenced, see Supporting Information, greatest skull height (GSH), taken from the lowest Table S1). Polymerase chain reaction amplification, point of the basioccipital to the highest point of the thermocycler settings and Sanger sequencing were cranium; postorbital width (POB), narrowest dorsal the same as those described by Demos et al. (2018) width posterior to the postorbital at the constriction and Patterson et al. (2018). Chromatograms were of the cranium; greatest mastoid breadth (MAST), edited and assembled in GENEIOUS PRO v.11.1.5 greatest breadth of cranium at mastoid processes; and (Biomatters Ltd). Sequence alignments were made greatest mandible length (MAND), taken from the using MUSCLE (Edgar, 2004) with default settings posteriormost point of the condyles to the anteriormost in GENEIOUS Prime v.2020.0.5. Protein-coding point of the incisors. The dental measurements sequence data from Cytb were translated to amino included: width across the third molars (M3–M3), taken acids to establish the absence of premature stop across the outermost point of the alveoli of the third codons, insertions and deletions. molars; complete upper canine–molar tooth row (C– The sequence alignments used in this study have M3), taken from the anteriormost point of the alveolus been deposited in the FIGSHARE data repository of the canine to the posteriormost point of the alveolus (https://doi.org/10.6084/m9.figshare.12698900.v1). of the third molar; width across upper canines (C–C), Newly generated sequence data have been deposited taken across the outermost points of the alveoli of the in GenBank under accession numbers (MT777844– canines; and complete mandibular canine–molar tooth

MT778066) (see also Supporting Information, Table S1). row (c–m3), taken from the anteriormost point of the

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 5 alveolus of the canine to the posteriormost point of We were unable to examine the type specimen of the third molar. Tooth abbreviations are as follows: C, Vespertilio hesperida Temminck, 1840 (= Pipistrellus Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 canine; I, incisor; M, molar; P, premolar; with upper hesperidus) [SMF 12381 (lectotype)], but this specimen teeth presented in upper case and lower teeth in has received detailed attention in the literature, lower case. including its history, type locality and detailed We examined type specimens (listed in Supporting description of characters together with craniodental Information, Table S1) from the following collections: measurements (Kock, 2001). The type locality is not The Natural History Museum (formerly The British definitely certain but is probably coastal Eritrea (Kock, Museum of Natural History), London (BMNH); the 2001). Other taxa that have been synonymized with Muséum national d’Histoire Naturelle, Paris (MNHN); Pip. hesperidus include Vesperugo subtilis Sundevall, Zoologisches Forschungsmuseum Alexander Koenig, 1846 (South Africa), Pipistrellus kuhlii fuscatus Bonn (ZFMK); the Durban Natural Science Museum, Thomas, 1901 (Kenya) and Pipistrellus (Romicia) Durban (DM); and The National Museum of Natural kuhli broomi Roberts, 1948 (South Africa). Whether History, Washington (USNM). any of these represent subspecies remains uncertain, The type specimens examined were as follows: although there appears to be little difference in the Vespertilio capensis A. Smith, 1829 [BMNH size of this species across its range (Kearney, 2013). 1849.8.16.21 (lectotype)], Vespertilio nanus Peters, Spatial relationships among the aforementioned 1852 [BMNH 1907.1.1.421 (syntype)], Neoromicia type specimens and other taxa of African and roseveari Monadjem et al., 2013 [DM 12617 Madagascarian Pipistrellini and Vespertilionini are (holotype)], Scotophilus rusticus Tomes, 1861 [BMNH shown in Figure 1. 1907.1.1.419 (lectotype)], Vesperus tenuipinnis Peters, In order to compare the morphology of the various 1872 [DM 13235 (neotype)], Vesperugo grandidieri taxa presented in this study, a principal components

Dobson, 1876 [MNHN 1996-2129 (holotype)], analysis (PCA) of log10-transformed values of Vesperugo (Vesperus) brunneus Thomas, 1880 [BMNH craniodental measurements (for a list of the most 1880.7.21.7 (holotype)], Vesperus bicolor Bocage, 1889 inclusive set of measurements available for each [BMNH 1889.5.1.3 (syntype)], Vesperugo (Vesperus) analysis, see Supporting Information, Tables S2 rendalli Thomas, 1889 [BMNH 1889.3.2.3 (holotype)], and S3) was conducted on the variance–covariance Vesperugo (Vesperus) flavescens Seabra, 1900 [MNHN matrix in the package ‘vegan’ (Oksanen et al., 2019) 1900-537 (syntype)], Vesperugo anchietae Seabra, 1900 run in R v.3.6.2 (R Core Team, 2019) and plotted [BMNH 1906.1.3.1 (syntype); we follow Kock (2001) using ggplot2 (Wickham, 2016). We first compared in accepting this as a justified emendation, because the skulls of the species traditionally placed in the Seabra (1900a) originally published the name as genus Neoromicia (e.g. Simmons, 2005) with the four ‘Vesperugo anchieta’ and immediately corrected it to recognized Laephotis species. We then compared the Vesperugo anchietae in Seabra (1900b)], Pipistrellus skulls of the ‘Neoromicia capensis’ group, including minusculus Miller, 1900 [USNM 84500 (holotype)], Neoromicia capensis (A. Smith, 1829) and Neoromicia Vespertilio minutus somalicus Thomas, 1901 [BMNH stanleyi Goodman et al., 2017 from the mainland and 1898.6.9.1 (holotype)], Pipistrellus crassulus Thomas, Neoromicia malagasyensis (Peterson et al., 1995), 1904 [BMNH 1904.2.8.1 (holotype)], Pipistrellus Neoromicia matroka Thomas & Schwann, 1905 nanulus Thomas, 1904 [BMNH 1904.2.8.8 (holotype)], and Neoromicia robertsi Goodman et al., 2012 from Pipistrellus helios Heller, 1912 [USNM 181813 Madagascar. For the analysis involving Neoromicia (holotype)], Pipistrellus aero Heller, 1912 [USNM and Laephotis, we used a subset of nine craniodental 181812 (holotype)], Pipistrellus musciculus Thomas, measurements, owing to missing measurements from 1913 [BMNH 1913.2.8.1 (holotype)], Eptesicus some specimens; these were as follows: GSKL, GSH, 3 3 3 ugandae Hollister, 1916 [USNM 166520 (holotype)], GBW, MAST, MAND, C–M , C–C, M –M and c–m3 (see Pipistrellus eisentrauti Hill, 1968 [BMNH 1967.2129 above for definitions of these terms). (paratype)], Laephotis botswanae Setzer, 1971 [USNM 425349 (holotype)], Laephotis namibensis Setzer, 1971 [USNM 342152 (holotype)], Neoromicia Bacular preparation isabella Decher, Hutterer & Monadjem, 2015 [ZFMK The baculum (os penis) of selected specimens was 2008.0292 (holotype)], Parahypsugo happoldorum prepared by severing the glans penis, rehydration in Hutterer et al., 2019 [ZFMK 2009.0029 (holotype)] water, then immersion in dilute sodium hydroxide that and Pipistrellus simandouensis Monadjem et al., 2020 was heated to 85 °C for a variable period. To facilitate [ZFMK 2008-0302 (holotype)]. Of these, the last three dissection, the glans was then stained in Alizarin Red, mentioned type specimens were sequenced; therefore, which is calcium specific and aids in distinguishing comparative genetic material was available for them. the bony baculum from investing tissues. Bacula

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 6 A. MONADJEM ET AL. Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020

Figure 1. Type localities of taxa of African and Malagasy Vespertilionini and Pipistrellini. Valid species are denoted by filled circles, subspecies and synonyms by open circles and species described herein by stars: 1, Pipistrellus abaensis J. A. Allen, 1917; 2, N[ycticejus]. adovanus Heuglin, 1877; 3, Pipistrellus aero Heller, 1912; 4, Vespertilio pipistrellus var. africanus Rüppell, 1842; 5, Nycticeius africanus G. M. Allen, 1911; 6, Scotoecus albigula Thomas, 1909; 7, Scoteinus schlieffeni albiventer Thomas & Wroughton, 1908; 8, Scotophilus albofuscus Thomas, 1890; 9, Vesperugo anchietae Seabra, 1900; 10, Laephotis angolensis Monard, 1935; 11, Eptesicus capensis angolensis Hill, 1937; 12, Pipistrellus ariel Thomas, 1904; 13, Scotoecus artinii De Beaux, 1923; 14, Eptesicus ater J. A. Allen, 1917; 15, Pipistrellus nanus australis Roberts, 1913; 16, Scoteinus schlieffeni australis Thomas & Wroughton, 1908; 17, Scoteinus schlieffeni bedouin Thomas & Wroughton, 1908; 18, Pipistrellus eisentrauti bellieri De Vree, 1972; 19, Hypsugo bemainty Goodman et al., 2015; 20, Vesperus bicolor Bocage, 1889; 21, Laephotis botswanae Setzer, 1971; 22, Pipistrellus (Romicia) kuhlii broomi Roberts, 1948; 23, Vesperugo (Vesperus) brunneus Thomas, 1880; 24, Vespertilio capensis A. Smith, 1829; 25, Scotoecus cinnamomeus Wettstein, 1916; 26, Pipistrellus

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 7 were photographed under a stereo microscope. Their (Afronycteris gen. nov., Hypsugo, Laephotis, Neoromica, morphological descriptions follow the convention of Nycticeinops and Pseudoromicia gen. nov.). Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 Hill & Harrison (1987). In the unique haplotype Cytb tree (Fig. 3), several genus-level clusters of taxa are apparent, but these do not coincide with current generic usage. The genus RESULTS Neoromicia as traditionally used is clearly paraphyletic. Some species (Neo. capensis, Neo. malagasyensis, Phylogenetic analyses Neo. matroka, Neo. robertsi, Neo. stanleyi and a new A preliminary alignment of 418 Cytb sequences was species from East Africa) assigned to that genus are assembled that included 310 sequences newly generated more closely related to the four Laephotis species for this study and 108 sequences downloaded from than to other Neoromicia. This group of ten species GenBank (Supporting Information, Table S1). Identical (plus Neo. cf. kirinyaga) is well supported as sister haplotypes were pruned from this alignment, resulting to another group that contains the type species of in a 333-sequence alignment (85% complete coverage) Neoromicia, Neoromicia zuluensis (Roberts, 1924), in that was used in ML and BI phylogenetic analyses close association with Neo. bemainty (Goodman et al., (the complete Cytb ML tree is shown in Supporting 2015), Neo. somalica (Thomas, 1901), Neo. cf. somalica Information, Fig. S1). The best-supported substitution and Neo. cf. guineensis (Fig. 3C). As here circumscribed, model estimated by jMODELTEST2 for the 333 bp an expanded Laephotis and a restricted Neoromicia Cytb alignment is GTR+I+G. Only the ML topology is are sister to another pair of genus-level clusters: one shown (Figs 2, 3), but both bootstrap (BS) values and comprising predominantly tropical rainforest species posterior probabilities (PPs) are depicted at shared, currently assigned to Neoromicia [Neo. brunnea well-supported nodes. Figure 2 depicts phylogenetic (Thomas, 1880), Neo. isabella, Neo. rendalli (Thomas, relationships among, and branch length ranges within, 1889) and Neo. roseveari], and this well-supported three genera of Pipistrellini (Pipistrellus, Scotoecus clade is sister to another distinctive group containing and Vansonia) and six genera of Vespertilionini only Neo. nana (Fig. 3B). These novel groupings are

crassulus Thomas, 1904; 27, Pipistrellus culex Thomas, 1911; 28, Vesperus damarensis Noack, 1889; 29, Scotophilus darwini Tomes, 1859; 30, Pipistrellus deserti Thomas, 1902; 31, Pipistrellus eisentrauti Hill, 1968; 32, Scotoecus falabae Thomas, 1915; 33, Eptesicus faradjius J. A. Allen, 1917; 34, Scoteinus schlieffeni fitzsimonsi Roberts, 1932; 35, Pipistrellus fouriei Thomas, 1926; 36, Pipistrellus kuhlii fuscatus Thomas, 1901; 37, Pipistrellus fuscipes Thomas, 1913; 38, Eptesicus garambae J. A. Allen, 1917; 39, Vespertilio capensis gracilior Thomas & Schwann, 1905; 40, Vesperugo (Vesperus) grandidieri Dobson, 1876; 41, Vesperus guineensis Bocage, 1889; 42, Pipistrellus hanaki Hulva & Benda, 2004; 43, Parahypsugo happoldorum Hutterer, Decher, Monadjem & Astrin, 2019; 44, Pipistrellus helios Heller, 1912; 45, Vespertilio hesperida Temminck, 1840; 46, Scotoecus hindei Thomas, 1901; 47, Scotophilus hirundo de Winton, 1899; 48, Vesperus humbloti Milne-Edwards, 1881; 49, Vesperugo hypoleucus Heuglin [in Fitzinger & Heuglin], 1866; 50, Pipistrellus inexspectatus Aellen, 1959; 51, Neoromicia isabella Decher, Hutterer & Monadjem, 2016; 52, Laephotis kirinyaga Monadjem et al., this paper; 53, Pseudoromicia kityoi Monadjem et al., this paper; 54, Hypsugo lanzai Benda, Al-Jumaily, Reiter & Nasher, 2011; 55, Pipistrellus leucomelas Monard, 1932; 56, Parahypsugo macrocephalus Hutterer & Kerbis Peterhans, 2019; 57, Vesperugo maderensis Dobson, 1878; 58, Eptesicus somalicus malagasyensis Peterson, Eger & Mitchell, 1995; 59, Vespertilio marginatus Cretzschmar, 1830; 60, Pipistrellus marrensis Thomas & Hinton, 1923; 61, Vespertilio matroka Thomas & Schwann, 1905; 62, Pipistrellus africanus meesteri Kock, 2001; 63, Eptesicus melckorum Roberts, 1919; 64, Scotophilus minimus Noack, 1887; 65, Pipistrellus minusculus Miller, 1900; 66, Vespertilio minuta Temminck, 1840; 67, Pipistrella minuta Loche, 1867; 68, Pipistrellus musciculus Thomas, 1913; 69, Laephotis namibensis Setzer, 1971; 70, Pipistrellus nanulus Thomas, 1904; 71, Vespertilio nanus Peters, 1852; 72, Eptesicus capensis nkatiensis Roberts, 1932; 73, Scabrifer notius G. M. Allen, 1908; 74, Pseudoromicia nyanza Monadjem et al., this paper; 75, †Scotoecus olduvensis Gunnell, Butler, Greenwood & Simmons, 2015; 76, Vesperugo pagenstecheri Noack, 1889; 77, Pipistrellus (Pipistrellus) permixtus Aellen, 1957; 78, Eptesicus phasma G. M. Allen, 1911; 79, Vespertilio pipistrellus Schreber, 1774; 80, Vespertilio platycephalus Temminck, 1832; 81, Vesperugo pulcher Dobson, 1875; 82, Vesperugo pusillulus Peters, 1870; 83, Pipistrellus raceyi Bates et al., 2006; 84, Eptesicus rectitragus Wettstein, 1916; 85, Vesperugo (Vesperus) rendalli Thomas, 1889; 86, Neoromicia robertsi Goodman et al., 2012; 87, Neoromicia roseveari Monadjem et al., 2013; 88, V[espertilio]. rueppelii J. Fischer, 1829; 89, Scotophilus rusticus Tomes, 1861; 90, Vespertilio savii Bonaparte, 1837; 91, Nycticejus schlieffenii Peters, 1859; 92, Pipistrellus rueppelli senegalensis Dorst, 1960; 93, †Nycticeinops serengetiensis Gunnell et al., 2015; 94, Pipistrellus simandouensis Monadjem et al., 2020; 95, Vespertilio minutus somalicus Thomas, 1901; 96, Vesperugo stampflii Jentink, 1888; 97, Neoromicia stanleyi Goodman et al., 2017; 98, Vesperus tenuipinnis Peters, 1872; 99, Eptesicus ugandae Hollister, 1916; 100, Neoromicia vansoni Roberts, 1932; 101, Pipistrellus vernayi Roberts, 1932; 102, Laephotis wintoni Thomas, 1901; 103, Scotoecus woodi Thomas, 1917; 104, Eptesicus zuluensis Roberts, 1924. Not mapped: [Pipistrellus Kuhli] latastei Laurent, 1937; Vespertilio pusillus LeConte, 1857; Vesperugo subtilis Sundevall, 1846.

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 8 A. MONADJEM ET AL. Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020

99/0.96 Laephotis

100/0.96

100/1.0 Neoromicia Ve spertilionini

100/1.0 Afronycteris gen. nov.

99/- 79/0.92

96/1.0 Pseudoromicia gen. nov.

90/0.99 Nycticeinops

96/1.0 97/1.0 100/1.0 Hypsugo Pipistrellin

85/0.93 63/0.98 Pipistrellus

70/0.68

96/1.0 i 78/0.91 Scotoecus 100/1.0 Vansonia 100/1.0 } outgroups

0.09

Figure 2. Maximum likelihood phylogeny of intergeneric relationships of mitochondrial cytochrome b sequences of Vespertilionidae. The phylogeny was inferred in IQ-TREE, and its topology was similar to the Bayesian phylogeny calculated in MrBayes. Bootstrap (BS) values followed by Bayesian posterior probabilities (PP) are indicated adjacent to nodes (those nodes with both BS < 70% and PP < 0.95 are not labelled).

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 9

to Fig. 3b. Tanzania FMNH208058

Tanzania FMNH208061 Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 7 Tanzania FMNH208059 7 Kenya Marsabit NMK184271 Kenya Marsabit NMK184170 Kenya Marsabit NMK184169 Kenya Laikipia NMK184515 Kenya Laikipia NMK184534 Kenya Laikipia NMK184605 Kenya Marsabit NMK184233 Kenya Marsabit NMK184286 Kenya Marsabit NMK184323 Kenya Marsabit NMK184268 Kenya Marsabit NMK184267 Kenya Marsabit NMK184325 Kenya Marsabit NMK184327 Pipistrellus hesperidus Kenya Laikipia NMK184431 Kenya Narok FMNH225628 Kenya Laikipia NMK184645 Kenya Narok FMNH225627 Malawi FMNH212006 Malawi FMNH212007 Madagascar KM886007 Mozambique JAG448 Mozambique JAG450 Mozambique JAG447 Madagascar KM886047 South Africa AJ841968 South Africa KM886097 Kenya Laikipia NMK184528 Kenya Nakuru FMNH216791 Kenya Laikipia NMK184481 Ethiopia KX375157 Guinea MK188529 Pipistrellus simandouensis Senegal JX276312 Senegal JX276313 Senegal JX276311 Pipistrellus cf. hesperidus Senegal JX276310 Senegal JX276314 Namibia KX375167 Pipistrellus rusticus Libya KM252757 Morocco KM252765 Morocco KM252761 Pipistrellus kuhlii Yemen KX375161 Guinea MK188530 Guinea ZWW273 Liberia ZWW063 Guinea ZWW272 Guinea ZWW276 Liberia ZWW044 Pipistrellus nanulus Tanzania FMNH192954 Tanzania FMNH219277 Tanzania FMNH193071 Uganda FMNH233016 Kenya Kakamega NMK184969 Madagascar KM886080 Madagascar KM886094 Pipistrellus raceyi Czech Republic AY582289 Morocco KM252778 Pipistrellus pipistrellus Portugal JX566929 Pipistrellus pygmaeus Sweden DQ120850 Kenya Isiolo FMNH220821 Pipistrellus nathusii Kenya Meru FMNH220828 Kenya Meru FMNH220816 Kenya Makueni FMNH221032 Scotoecus hindei Kenya Meru FMNH216110 Kenya Narok FMNH225594 Kenya Narok FMNH225595 Senegal JX276317 Scotoecus hirundo Kenya Kisumu FMNH215624 Kenya Kisumu NMK184939 Rwanda FMNH225229 Vansonia cf. rueppellii Rwanda FMNH225250 Egypt KX375170 Senegal JX276316 Vansonia rueppellii Mimetillus moloneyi Kerivoula argentata Scotophilus nigrita Glauconycterus variegata Myotis tricolor Miniopterus fraterculus

0.09

Figure 3. Maximum likelihood phylogeny of mitochondrial cytochrome b sequences of Vespertilionidae: (A) Pipistrellus, Scotoecus, Vansonia, and outgroups (B) Afronycteris, Pseudoromicia, Nycticeinops, and Hypsugo (C) Laephotis and Neoromicia. The phylogeny was inferred in IQ-TREE, and its topology was similar to the Bayesian phylogeny calculated in MrBayes. Filled red circles on nodes denote bootstrap (BS) values ≥ 70% and Bayesian posterior probabilities (PP) ≥ 0.95. Open circles outlined in black indicate BS ≥ 70% and PP < 0.95, and open circles outlined in red indicate BS < 70% and PP > 0.95. Support values for most minor clades are not shown. Specimen localities include counties for Kenya. DRC refers to Democratic Republic of the Congo and CAR to Central African Republic. Museum acronyms are defined in the Material and Methods section. Sequences downloaded from GenBank are indicated by inclusion of GenBank accession numbers (Supporting Information, Table S1). Branch colours indicate individual species/clade membership.

described below as the new genera Pseudoromicia and paraphyletic. The group also includes Pip. grandidieri, Afronycteris, respectively. The remaining sub-Saharan the type species of subgenus Afropipistrellus (Fig. 3B). Vespertilionini comprise two clusters: the genus This expanded group of Nycticeinops lineages is sister Nycticeinops (type species Nycticeinops schlieffeni to a largely Palaearctic and Indo-Malayan cluster of Peters, 1859), hitherto considered monospecific, is Hypsugo species. flanked by species assigned to Parahypsugo (type In contrast to the novel groupings found for species Par. happoldorum), rendering that taxon Vespertilionini, our analyses of Pipistrellini confirmed

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 10 A. MONADJEM ET AL.

to Fig. 3c

Kenya Kisumu FMNH215615 Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 Kenya Kisumu FMNH215616 Kenya Kisumu FMNH234689 Kenya Kisumu FMNH234588 Guinea ZWW203 Guinea ZWW202 Uganda FMNH233018 Rwanda FMNH225212 Uganda FMNH160366 Rwanda FMNH225231 Uganda FMNH160365 Uganda FMNH160375 DRC FMNH173300 Mozambique FMNH177148 Tanzania FMNH205243 Malawi FMNH192110 Uganda FMNH233034 Uganda FMNH233037 Mozambique FMNH177216 Tanzania FMNH187144 Tanzania FMNH187225 Mozambique FMNH177218 Tanzania FMNH187224 Tanzania FMNH192683 Botswana KM886074 Zambia KX375181 Botswana KM886076 Namibia KX375180 Tanzania FMNH151202 Tanzania FMNH192902 Afronycteris nana DRC FMNH173297 DRC FMNH173301 DRC FMNH173299 Rwanda FMNH225230 Mozambique FMNH177149 Mozambique FMNH177217 Zambia KX375182 Ethiopia EU797428 Senegal JX276107 Senegal JX276195 Senegal JX276178 Senegal JX276203 Senegal JX276160 Kenya Meru FMNH220923 Kenya Meru FMNH220927 Kenya Samburu FMNH220947 Kenya Meru FMNH220926 Kenya Kitui FMNH234716 Kenya Kitui FMNH234715 Kenya Kitui NMK185098 Kenya Meru FMNH220911 Kenya Kitui FMNH234592 Kenya Kilifi FMNH234597 Tanzania FMNH187391 Tanzania FMNH187390 Kenya Narok FMNH225618 Kenya Narok FMNH225622 Kenya Narok FMNH225621 Kenya Kisumu FMNH215628 Kenya Kisumu FMNH215629 Kenya Homa Bay FMNH215658 Kenya Kisumu NMK184934 Kenya Kisumu NMK184940 Pseudoromicia nyanza sp. nov. Kenya Kisumu NMK184938 Kenya Homa Bay FMNH215661 Kenya Homa Bay FMNH215659 Kenya Narok FMNH225619 Kenya Narok FMNH225617 Kenya Narok FMNH225620 Guinea MK188528 Guinea ZWW263 Liberia ZWW045 Liberia ZWW048 Pseudoromicia roseveari Guinea ZWW251 Liberia ZWW017 Uganda FMNH223211 Uganda FMNH223555 Pseudoromicia kityoi sp. nov. Tanzania FMNH192956 Pseudoromicia sp. Guinea MK188527 Pseudoromicia isabella Guinea ZWW175 Guinea ZWW282 Pseudoromicia brunnea Guinea ZWW136 Senegal JX276206 Senegal JX276207 Central African Republic JQ956446 Pseudoromicia rendalli Kenya Malindi FMNH216159 Kenya Malindi FMNH216119 Kenya Meru FMNH216109 Kenya Kilifi FMNH216112 Kenya Kilifi FMNH216113 Kenya Kilifi FMNH216114 Kenya Kilifi FMNH216115 Kenya Malindi FMNH216118 Nycticeinops grandidieri Kenya Kilifi FMNH234596 Kenya Kakamega FMNH234598 Mozambique JAG430 Mozambique JAG439 Mozambique JAG441 Mozambique JAG440 Guinea MK188525 Guinea ZWW275 Guinea MK188523 Guinea MK188524 Nycticeinops happoldorum Guinea MK188526 Guinea MK188520 Guinea MK188521 Nycticeinops bellieri Tanzania FMNH192955 Nycticeinops cf. crassulus Senegal JX276302 Senegal JX276308 Senegal JX276303 Nycticeinops schlieffeni Senegal JX276305 Senegal JX276301 Kenya Kajiado FMNH221014 Kenya Taita Taveta FMNH221015 Nycticeinops cf. schlieffeni Kenya Kitui FMNH234424 Cameroon MK188522 Nycticeinops eisentrauti Oman KX375190 Hypsugo arabicus Greece DQ120866 Hypsugo savii Vietnam KX429687 Hypsugo dolichodon Egypt KX375193 Hypsugo ariel to Fig. 3a Laos DQ318883 Hypsugo cadomae Malaysia KX429685 Hypsugo macrotis Laos KX429686 Hypsugo pulveratus

Figure 3. Continued.

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 11

Tanzania FMNH168093 Tanzania FMNH168094

Tanzania FMNH219271 Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 Tanzania FMNH187221 Tanzania FMNH219120 Tanzania FMNH219275 Tanzania FMNH219111 Tanzania FMNH219270 Tanzania FMNH219274 Tanzania FMNH219113 Laephotis capensis Tanzania FMNH219266 Botswana KM886073 South Africa KX375174 Malawi FMNH212008 Tanzania FMNH219114 Namibia KX375173 Zambia KX375175 South Africa FMNH195628 South Africa KM886041 South Africa FMNH195629 Madagascar FMNH222777 Madagascar KM886037 Madagascar FMNH222778 Madagascar FMNH222779 Madagascar FMNH222785 Laephotis matroka Madagascar FMNH222781 Madagascar FMNH222782 Madagascar FMNH222784 Madagascar Madagascar KM886006 Kenya Marsabit NMK184174 Kenya Marsabit NMK184400 Ethiopia KX375185 Kenya Marsabit NMK184337 Kenya Marsabit NMK184399 Kenya Kakamega NMK184868 Kenya Marsabit NMK184265 Kenya Laikipia NMK184583 Laephotis kirinyaga sp. nov. Kenya Laikipia NMK184665 Uganda FMNH233035 Uganda FMNH233036 Ethiopia KX375184 Ethiopia KX375183 Senegal JX276106 Tanzania FMNH219268 Tanzania FMNH219273 Laephotis cf. kirinyaga Kenya Samburu FMNH220896 Kenya Samburu FMNH220897 Kenya Laikipia NMK184627 Laephotis wintoni Kenya Laikipia NMK184523 Kenya EU797434 Namibia KX375171 Zambia KX375172 South Africa EU797444 Laephotis botswanae South Africa EU797445 Namibia EU797438 Namibia EU797443 Laephotis namibensis South Africa EU797435 South Africa EU797446 Laephotis cf. wintoni Madagascar KM886030 Madagascar KM886078 Madagascar FMNH213931 Laephotis robertsi Madagascar FMNH175988 Madagascar KM886025 Laephotis malagasyensis Botswana KM886072 Botswana KM886077 Tanzania FMNH187143 Laephotis stanleyi Tanzania FMNH187223 Tanzania FMNH219264 Tanzania FMNH219278 Malawi FMNH211514 Kenya Narok FMNH225614 Kenya Narok FMNH225615 Kenya Kajiado FMNH221026 Kenya Narok FMNH225613 Kenya Samburu FMNH221025 Kenya Kajiado FMNH221017 Kenya Narok FMNH225623 Kenya Narok FMNH225624 Kenya Narok FMNH225616 Kenya Narok FMNH225612 Neoromicia somalica Kenya Narok FMNH225625 Kenya Kajiado FMNH221016 Kenya Kajiado FMNH221018 Kenya Laikipia FMNH234671 Kenya Laikipia FMNH234679 Kenya Laikipia FMNH234677 Kenya Kakamega FMNH234585 Kenya Kakamega FMNH234587 Kenya Kakamega FMNH215614 Kenya Narok FMNH225820 Yemen KX375178 Yemen KX375179 Neoromicia cf. guineensis Yemen KX375177 Kenya Makueni FMNH221022 Kenya Taita Taveta FMNH221023 Kenya Samburu FMNH220910 Kenya Samburu FMNH220913 Kenya Samburu FMNH220919 Kenya Samburu FMNH220936 Kenya Samburu FMNH220906 Kenya Kitui FMNH234600 Kenya Kitui FMNH234691 Kenya Kitui NMK185143 Kenya Samburu FMNH220943 Kenya Kitui NMK185107 Kenya Samburu FMNH220914 Kenya Samburu FMNH220937 Neoromicia zuluensis Kenya Samburu FMNH220918 Kenya Samburu FMNH220940 Kenya Kitui NMK185115 Kenya Kitui NMK185132 Kenya Kilifi FMNH216160 Kenya Kilifi FMNH216161 South Africa FMNH195630 South Africa FMNH195631 South Africa KX375186 South Africa FMNH195632 Namibia KX375187 Namibia KX375188 Senegal JX276262 Senegal JX276290 Senegal JX276218 Senegal JX276208 Senegal JX276230 Senegal JX276300 Neoromicia cf. somalica Senegal JX276211 Senegal JX276248 Senegal JX276215 Senegal JX276232 Madagascar KM886043 Neoromicia bemainty to Fig. 3b.

Figure 3. Continued.

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 12 A. MONADJEM ET AL. the existing content and composition of the three and low loadings and reflects differences in shape. genera found in the Afrotropics: our sample included The largest positive loading is with GSKL (0.488) Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 ten species of Pipistrellus, which are sister to the and the largest negative loading with GSH (−0.656), species of Scotoecus; this group is then sister to suggesting that species with higher projections on PC2 Vansonia, comprising only V. rueppellii (Fig. 3A). have larger, flatter crania than those with lower ones. The genetic distances for the various genera are These four groups neatly correspond to four distinct shown in Table 1, and distances for 53 Pipistrellini bacular types (Fig. 5), and we suggest that they and Vespertilionini species are shown in the represent different genera (Taxonomic conclusions, see Supporting Information (Table S4). It is noteworthy below). The bacula of these four groups differ in size, that genetic distances for the two newly identified the shape of the shaft and the shape of the proximal genera are comparable to other long-recognized and distal ends (see Fig. 5). The baculum shape of each genera. The genetic distances among the seven group is described in detail in the description of the currently recognized genera in this study (Hypsugo, genera (see below). Laephotis, Neoromicia, Nycticeinops, Pipistrellus, Within the ‘Neo. capensis’ group, the PCA ordination Scotoecus and Vansonia) range from 0.148 to 0.203 on craniodental measurements shows that most (with intrageneric distances ranging from 0.042 to species occupy separate regions of morphospace 0.142). In comparison, the new genera Pseudoromicia (Fig. 6). The first two principal axes account for > 81% and Afronycteris have genetic distances ranging of the variation, with the first axis representing a size from 0.157 to 0.196 (with intrageneric distances of gradient with negative loadings on all measurements 0.097 and 0.032, respectively). Afronycteris nana is (Supporting Information, Table S3). Hence, the largest well supported as monophyletic, as are five of seven species (e.g. Neo. stanleyi and Neo. robertsi) are on species within Pseudoromicia; the exceptions being the left of the ordination and the smallest species Pseudoromicia isabella and Pse. sp. from Tanzania (e.g. Neo. matroka) on the right. However, there is represented by single sequences on relatively long significant overlap between Neo. matroka and Neo. branches (Fig. 3B). cf. capensis. The second principal component has both high and low loadings and represents differences in shape, with all long axis length measurements having Morphometric analyses positive values and all width measurements (except The PCA ordination on craniodental measurements MAST and C–C) having negative values (Supporting shows that the species traditionally placed within Information, Table S3). The largest positive loading

Neoromicia and Laephotis fall into four distinct is with c–m3 (0.157) and the largest negative loading regions of morphospace (Fig. 4). The first two principal with POB (−0.727), suggesting that species with high axes account for > 92% of the variation, with the positive values on PC2 have longer mandibles and first axis (PC1) representing a size gradient with narrower post-orbital constrictions compared with negative loadings on all measurements (Supporting those having lower values. Furthermore, Neo. capensis Information, Table S2). Hence, the largest species from southern Africa occupies a mostly different (e.g. Neo. robertsi) appear on the left of the ordination morphospace compared with Neo. cf. capensis from and the smallest species (e.g. Neo. nana) on the right. East and West Africa, which we describe as a new The second principal component (PC2) has both high species.

Table 1. Uncorrected cytochrome b p-distances among (below diagonal) and within (numbers on diagonal, in bold) nine genera of Vespertilionidae calculated in MEGA X v.10.0.5

Genus 1 2 3 4 5 6 7 8 9

1 Afronycteris gen. nov. 0.032 2 Hypsugo 0.189 0.142 3 Laephotis 0.169 0.174 0.076 4 Neoromicia 0.183 0.181 0.148 0.069 5 Nycticeinops 0.187 0.183 0.172 0.178 0.121 6 Pipistrellus 0.196 0.198 0.185 0.182 0.188 0.107 7 Pseudoromicia gen. nov. 0.168 0.181 0.157 0.165 0.178 0.182 0.097 8 Scotoecus 0.190 0.198 0.176 0.197 0.178 0.175 0.181 0.058 9 Vansonia 0.196 0.200 0.190 0.203 0.199 0.192 0.189 0.188 0.042

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Figure 4. Principal components analysis of craniodental measurements of the species traditionally allocated to the genus Neoromicia s.l.; the colours correspond to the four distinct types of bacula exhibited by these species (shown in Fig. 5). Each dot refers to the mean value of the craniodental measurements of examined specimens; see main text for further details.

Taxonomic conclusions Furthermore, we suggest that the ‘Laephotis’ Our phylogenetic analyses show support for the genera clade (Fig. 2), which includes the genus Laephotis Pipistrellus and Vansonia, but clearly indicate the and several species currently placed in the genus paraphyly of the genera Neoromicia and Parahypsugo Neoromicia (including Neo. capensis, Neo. stanleyi as currently recognized (Moratelli & Burgin, 2019; and the Malagasy species Neo. matroka, Neo. robertsi Monadjem et al., 2020a). and Neo. malagasyensis), be recognized under the Our phylogeny supports synonymizing genus Laephotis. The third clade, which includes Parahypsugo with Afropipistrellus, because the mostly tropical rainforest species of Neoromicia s.l. addition of Nyc. grandidieri to the group of species (including Neo. brunnea, Neo. roseveari, Neo. isabella recognized by Hutterer et al. (2019) as Parahypsugo and Neo. tenuipinnis), in addition to the widely [type species Par. happoldorum plus Par. bellieri (De distributed Neo. rendalli, currently has no pre- Vree, 1972), Par. crassulus (Thomas, 1904) and Par. existing name, for which we describe a new genus (see eisentrauti (Hill, 1968), with Par. macrocephalus below). A new genus is also needed for the ubiquitous added subsequently by Hutterer & Kerbis Peterhans banana bat, formerly known as Neo. nana. (2019)] renders Parahysugo a junior synonym of Our phylogeny supports the recognition of Afropipistrellus. The strong divergence of ‘Par. Vansonia as a distinct genus (rather than a subgenus eisentrauti’ places it immediately outside the group of Pipistrellus), because it is sister to Pipistrellus + Afropipistrellus + Nycticeinops, but this appraisal Scotoecus. This generic rearrangement of the African is based on only 764 bp of Cytb. Therefore, we members of the Vespertilionini and Pipistrellini is also propose that Afropipistrellus and Parahypsugo be reflected in certain morphological traits, particularly synonymized with Nycticeinops. bacular shape and, where information is available, We suggest restricting Neoromicia to the type penial characteristics (Fasel et al., in press). Based on species Neo. zuluensis, the sister species Neo. somalica the molecular and morphological evidence presented and the sister taxa Neo. bemainty and Neo. anchietae. above, we also describe three new species.

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 14 A. MONADJEM ET AL.

Description: This genus was originally created for the species Neo. zuluensis, based on it having ‘the cranium Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 slightly raised above the level of the muzzle’ (Roberts, 1926). The close relationship between this taxon and Neo. somalica (Thomas, 1901) has long been recognized, and the two are sister taxa in our phylogeny. Based on our genetic and morphometric analyses presented above, we have expanded this genus further to include the following species: Neo. guineensis (Bocage, 1889), Neo. anchietae (Seabra, 1900) and Neo. bemainty (Goodman et al., 2015). These are all small-sized pipistrelle-like bats with a distinct bacular morphology (Fig. 5B). The baculum (~1.5–2.0 mm in length) is shorter than in Pseudoromicia and similar in length to that of Laephotis and Afronycteris. It has a characteristic shape, with a thick base that is weakly bilobed, a shaft with a straight outer margin and slightly curved inner margin, and a unique three-pronged (cross-shaped) tip that is set at a slight angle to the shaft (Fig. 5B). They also have a more inflated braincase than Laephotis, but not as inflated as Afronycteris, from which they differ in many other respects (for more details, see the description in the account of Afronycteris). They lack the white wings of Pseudoromicia and have bicoloured fur on both the upper and under parts. All five species are essentially savanna or woodland species, with four occurring in southern and eastern Africa and Madagascar.

Figure 5. Bacula of the four clades within formerly or traditionally recognized as Neoromicia: A, Laephotis Laephotis Thomas, 1901 kirinyaga (FMNH 234639); B, Neoromicia somalica (FMNH 215614); C, Pseudoromicia kityoi (FMNH 223211); and D, Synonymy Afronycteris nana (DM 13013). Note the three-pronged tip in Vespertilio A. Smith, 1829 (part, not Linnaeus, 1758). Neoromicia, the straight shaft with spatulate tip at an angle Hypsugo Kolenati, 1860 (part, not Kolenati, 1856). of 45° in Laephotis, the long, curved shaft with bilobed tip Scotophilus Thomas, 1861 (part, not Leach, 1821). in Pseudoromicia and the deeply bilobed base and gently Vesperugo Dobson, 1878 (part, not Keyserling & curved shaft in Afronycteris. Scale bars: 1 mm. Blasius, 1839). Vesperus Jentink, 1887 (part, not Keyserling & TAXONOMY Blasius, 1839). Family Vespertilionidae Gray, 1821 Eptesicus Matschie, 1897 (part, not Rafinesque, 1820). Scabrifer G.M. Allen, 1908. Tribe Vespertilionini Gray, 1821 Rhinopterus G.M. Allen, 1939 (part, not Miller, 1906). Neoromicia Roberts, 1926 Pipistrellus Heller & Volleth, 1984 (part, not Synonymy Kaup, 1829). Nycterikaupius (part, not Menu, 1987). Vesperugo Bocage, 1889 (part, not Keyserling & Neoromicia Volleth et al., 2001 (part, not Roberts, Blasius, 1839). 1926). Vespertilio Thomas, 1901 (part, not Linnaeus, 1758). Complete synonymic histories for the species of Eptesicus G.M. Allen, 1911 (part, not Laephotis are given in the African Chiroptera report Rafinesque, 1820). (AfricanBats NPC, 2019). Pipistrellus Zammarano, 1930 (part, not Kaup, 1829). Complete synonymic histories for the species of Neoromicia are given in the African Chiroptera report Description: This genus was originally created for the (AfricanBats NPC, 2019). species Laephotis wintoni Thomas, 1901, with the name

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Figure 6. Principal components analysis of craniodental characters of the short-eared species of Laephotis as recognized in this study. referring to the large ‘sail-like’ ears of that species. Laephotis kirinyaga Monadjem, Patterson, A second, closely related species with large ears was Webala & Demos sp. nov. described a quarter of a century later, Lae. angolensis East African serotine Monard 1935, and two more species by Setzer in 1971: LSID: http://zoobank.org/urn:lsid:zoobank.org:pub: Lae. botswanae and Lae. namibensis. The baculum 71737F08-2938-4403-8385-5438B2E5EABE (1.5–2.0 mm in length) of Laephotis as defined herein is shorter than in Pseudoromicia and similar in Synonymy length to that of Neoromicia and Afronycteris. It has a Eptesicus capensis Kingdon (1974). characteristic shape, with a bilobed base, straight shaft Pipistrellus capensis garambe Thorn & Kerbis and a spatulate tip that is at an angle of ~45° to the Peterhans (2009) (in part). shaft (Fig. 5A). Neoromicia capensis Patterson & Webala (2012). Based on our genetic and morphometric analyses Neoromicia somalica Benda et al. (2016) (in part). presented above, we have expanded further this genus to include the following species: Lae. capensis Holotype: FMNH 234558, field number BDP 7516. (A. Smith, 1829), Lae. matroka (Thomas & Schwann, This specimen was collected by Bruce D. Patterson, 1905), Lae. robertsi (Goodman et al., 2012), Lae. Paul W. Webala, Carl W. Dick and Beryl Makori. It is malagasyensis (Peterson et al., 1995) and Lae. stanleyi an adult male, with muscle tissue in liquid nitrogen, (Goodman et al., 2017). the body fixed in formalin and preserved in ethanol, Laephotis is readily distinguished by its bacular now with skull extracted and cleaned. morphology (Hill & Harrison, 1987). It is easily separated from Afronycteris based on external features (for details, see the account of Afronycteris). Type locality: Marsabit National Park, 1.3 km SE of This genus may also be distinguished from Neoromicia campground near Headquarters, Marsabit County, by its larger size. Furthermore, the cranium is more Kenya (2.3090°N, 38.0001°E; Fig. 1). The type specimen robust in Laephotis and obviously flattened compared was netted on 27 July 2015 at an elevation of 1280 m with Neoromicia and Pseudoromicia. Laephotis also above sea level. lacks the white wings of Pseudoromicia and is mostly associated with arid savannas and grasslands. Of the Paratypes: One other male (FMNH 234559) was nine species that we recognize in this genus, all except captured at the same location and on the same night as the one we describe here are restricted to eastern the type specimen and is considered a paratype. Seven and southern Africa and Madagascar, and none is other individuals (FMNH 234546, FMNH 234549– associated with rainforests of tropical Africa. 234553, FMNH 234556–234557, four males and three

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 16 A. MONADJEM ET AL. females), were collected close to the type locality at elevations ranging from 1157 to 1356 m from 16 to 26 Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 July 2015 (Supporting Information, Table S1); they closely resemble the holotype genetically (Fig. 3C) and morphologically (Tables 2–4) and are also considered 0.53, 4.5 ± 0.53, N = 4 3.7–6.0, 0.88, 5.3 ± 0.88, N = 12 4.0–7.5, 1.72, 8.5 ± 1.72, N = 5 7.4–11.5, 0.30, 6.5 ± 0.30, N = 7 6.1–6.8, 1.23, 3–8, 3–8, 5.7 ± 1.23, N = 18 0.87, 4–7, 4–7, 5.3 ± 0.87, N = 16 4.7 paratypes. Body mass

Etymology: The specific epithet is a Kikuyu word for Mount Kenya and reflects the distribution of the species in the northern highlands of Kenya. It is a noun in apposition.

Diagnosis: This species is similar in size and 0.96, 31–32, 31–32, 31.3 ± 0.96, N = 4 1.51, 30–34, 30–34, 32.1 ± 1.51, N = 12 1.41, 34.5–38.0, 34.5–38.0, 35.5 ± 1.41, N = 5 1.58, 34–39, 34–39, 37.0 ± 1.58, N = 8 2.22, 29.1–37.1, 29.1–37.1, 33.4 ± 2.22, N = 38 1.35, 30.5–34.1, 30.5–34.1, 31.5 ± 1.35, N = 16 31.5 appearance to its sister species Lae. capensis. It is length Forearm easily distinguished from the long-eared Laephotis species by its shorter ears. Of the short-eared Laephotis species, Lae. stanleyi and Lae. robertsi are significantly larger in forearm length and most craniodental measurements (Tables 2–4). In contrast, Lae. malagasyensis is smaller, especially in cranial measurements (Table 3). Laephotis matroka is similar in external and craniodental measurements 0.82, 11–13, 11–13, 12.0 ± 0.82, N = 4 0.76, 11–13, 11–13, 11.8 ± 0.76, N = 12 0, 13–13, N = 5 13–13, 13.0 ± 0, 1.91, 10–15, 10–15, 12.3 ± 1.91, N = 8 1.46, 11–17, 11–17, 12.8 ± 1.46, N = 24 1.10, 9–13, 9–13, 11.5 ± 1.10, N = 16 12 Ear length but is typically darker brown above and medium brown below (Goodman, 2011). In any case, Lae. robertsi, Lae. malagasyensis and Lae. matroka are all endemic to Madagascar (Goodman et al., 2012, 2017) and genetically distinct from Lae. kirinyaga (Fig. 3C). Laephotis kirinyaga closely resembles Lae. capensis, from which it differs by 8.3% on the Cytb 0.50, 4–5, 4–5, 4.8 ± 0.50, N = 4 0.51, 4–5, 4–5, 4.6 ± 0.51, N = 12 0.55, 5–6, 5–6, 5.4 ± 0.55, N = 5 0.76, 6–9, 6–9, 7.3 ± 0.76, N = 8 0.84, 6–8, 6–8, 7.4 ± 0.84, N = 24 0.97, 5–8, 5–8, 6.5 ± 0.97, N = 16 6 Hindfoot length gene (Supporting Information, Table S4). Externally, the two species are alike and broadly overlap in size, but Lae. kirinyaga is on average smaller in most measurements, particularly total length and forearm length (Table 2). Likewise, Lae. capensis is on average larger for all craniodental measurements (but with significant overlap), except greatest skull height, 1.00, 35–37, 35–37, 36.0 ± 1.00, N = 3 2.76, 27–36, 27–36, 32.2 ± 2.76, N = 12 2.07, 31–36, 31–36, 34.6 ± 2.07, N = 5 4.46, 34–48, 34–48, 39.7 ± 4.46, N = 5 3.83, 26–40, 26–40, 34.9 ± 3.83, N = 24 2.22, 28–37, 28–37, 32.6 ± 2.22, N = 16 32 which is greater in Lae. kirinyaga. This is borne out in length Tail the lateral profile of the skull (Fig. 7), which is visibly flatter in Lae. capensis. These two species occupy mostly separate regions in multivariate space (Fig. 6), but again with some overlap. In contrast, the three specimens assigned to Lae. kirinyaga from Ethiopia and Guinea (labelled ‘cf. capensisW’ in Fig. 6), for which genetic data are lacking, fall completely within 1.15, 80–82, 80–82, 81.3 ± 1.15, N = 3 3.18, 77–86, 77–86, 82.1 ± 3.18, N = 12 4.06, 84–94, 84–94, 91.0 ± 4.06, N = 5 6.42, 83–101, 83–101, 93.3 ± 6.42, N = 8 7.09, 76–102, 76–102, 88.7 ± 7.09, N = 24 5.67, 74–95, 74–95, 82.8 ± 5.67, N = 16 80 the multivariate space of the Lae. kirinyaga specimens length Total (labelled ‘cf. capensis’) that have been sequenced.

Description: Exernal characters: Laephotis kirinyaga is a medium-sized pipistrelle-like bat, with strongly contrasting fur dorsally and ventrally. The dorsal pelage is medium brown, with most individual hairs being tipped light yellowish brown, giving the bat a from Marsabit National Park, Kenya from Marsabit National Park, kirinyaga External measurements (in millimetres) and mass grams) of Laephotis brightly coloured appearance. The ventral pelage is cream–white to light cream–brown, with a dark base. (other specimens) The ears are short and rounded, and the tragus is Holotype FMNH 234558 Laephotis malagasyensis Laephotis matroka Laephotis robertsi Laephotis stanleyi Laephotis capensis kirinyaga Laephotis kirinyaga Laephotis ). Measurements are of the holotype, other individuals of the new species and other ‘short-eared’ species of Laephotis . other individuals of the new species and Measurements are of the holotype, range and sample size ( N ). Measurements are presented as the mean ± SD, are taken from Goodman et al. (2017) . malagasyensis , matroka and Lae. robertsi , Lae. Lae. Measurements for the three Malagasy endemics, curved distally on both anterior and posterior margins, 2. Table Specimen or taxon

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 17 Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 MAND 9.49 9.85 ± 0.33, 9.36–10.52, N = 16 10.35 ± 0.43, 9.41–11.17, N = 57 10.6 ± 0.32, 10.2–11.0, N = 6 10.6 ± 0.15, 10.4–10.7, N = 4 9.0 ± 0.21, 8.7–9.3, N = 10 8.3 ± 0.13, 8.2–8.5, N = 5 GSH 4.60 4.68 ± 0.30, 4.22–5.49, N = 16 4.57 ± 0.19, 4.17–5.11, N = 57 – – – – GBW 6.74 7.07 ± 0.22, 6.72–7.49, N = 16 7.28 ± 0.24, 6.81–7.72, N = 57 – – – – MAST 7.00 7.49 ± 0.32, 6.96–8.09, N = 16 7.87 ± 0.34, 7.07–8.74, N = 57 8.4 ± 0.10, 8.2–8.5, N = 7 8.4 ± 0.22, 8.2–8.7, N = 4 7.5 ± 0.25, 7.2–7.9, N = 12 7.2 ± 0.32, 6.9–7.7, N = 5 POB 3.62 3.71 ± 0.21, 3.34–4.16, N = 16 3.68 ± 0.13, 3.42–4.00, N = 57 3.7 ± 0.21, 3.5–4.1, N = 7 4.0 ± 0.24, 3.6–4.1, N = 4 3.5 ± 0.14, 3.3–3.8, N = 12 3.4 ± 0.18, 3.2–3.7, N = 5 ZYGO 8.95 ± 0.37, 8.24–9.62, N = 57 9.6 ± 0.29, 9.3–10.0, N = 6 9.9 ± 0.37, 9.5–10.4, N = 4 8.8 ± 0.28, 8.4–9.2, N = 9 8.5 ± 0.26, 8.3–8.8, N = 5 8.41 8.71 ± 0.37, 7.95–9.73, N = 16 GSKL 14.05 ± 0.48, 13.09–15.35, N = 58 15.2 ± 0.21, 14.8–15.4, N = 7 14.7 ± 0.19, 14.4–14.8, N = 4 13.1 ± 0.33, 12.4–13.4, N = 12 12.6 ± 0.38, 12.4–13.3, N = 5 13.21 13.55 ± 0.43, 12.89–14.25, N = 16 from Marsabit National Park, Kenya from Marsabit National Park, kirinyaga Cranial measurements (in millimetres) of specimens Laephotis

Holotype FMNH 234558 (other specimens) ). Measurements are of the holotype, other individuals of the new species and other ‘short-eared’ species of Laephotis . other individuals of the new species and Measurements are of the holotype, ( N ). range and sample size are presented as the mean ± SD, Measurements are taken from Goodman et al. (2017) . malagasyensis , and Lae. matroka robertsi , Lae. Lae. Measurements for the three Malagasy endemics, Table 3. Table Specimen or taxon kirinyaga Laephotis kirinyaga Laephotis Laephotis capensis Laephotis stanleyi Laephotis robertsi Laephotis matroka Laephotis malagasyensis

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Figure 7. Plate showing the cranium and mandible of Laephotis kirinyaga (FMNH 234558). Scale bar = 5 mm.

ending in a rounded tip, as in Lae. capensis (Monadjem et al., 2020b). The ears and muzzle are dark brown in colour, and the skin around the eyes is dark brown in the type specimen (Fig. 8A) but mostly pinkish in the paratypes. 4.13 0.18, 4.03–4.55, N = 16 4.03–4.55, 4.28 ± 0.18, 0.17, 4.02–4.81, N = 57 4.02–4.81, 4.45 ± 0.17, 0.20 4.3–4.8, N = 7 4.6 ± 0.20 4.3–4.8, N = 4 4.9 ± 0.17 4.6–5.0, N = 12 4.1 ± 0.17 3.9–4.4, N = 5 3.8 ± 0.16 3.6–4.0, C–C

Craniodental characters: The skull is relatively robust, as in Lae. capensis, but less so than in Lae. stanleyi. In lateral profile, the cranium is distinctly straight, rising only gently up from the rostrum to the top of the braincase. An occipital ‘helmet’ is present but poorly developed, and the sagittal and lambdoidal crests are visible. The zygomatic arches are relatively robust (Fig. 7), as in Lae. capensis. The dentition in

3 Lae. kirinyaga is typical of the genus, with I 2/3, C 1/1, P 1/3, M 2/3. In the upper tooth row, I1 is unicuspid C–M 4.75 0.17, 4.50–5.09, N = 16 4.50–5.09, 4.79 ± 0.17, 0.21, 4.49–5.53, N = 10 4.49–5.53, 4.96 ± 0.21, 0.13 4.9–5.3, N = 7 5.1 ± 0.13 4.9–5.3, N = 4 5.4 ± 0.13 5.2–5.5, N = 12 4.5 ± 0.17 4.2–4.8, N = 5 4.2 ± 0.13 4.1–4.4, 2 and I is small, not reaching halfway up the length of I1. The P1 is absent, putting C in contact with P2. The

m3 is myotodont sensu Van Cakenberghe & Happold (2013).

Biology: This species has been captured infrequently across the highlands of Kenya on both sides of the Rift Valley. It is present in wet tropical forest (e.g. Kakamega from Marsabit National Park, Kenya from Marsabit National Park, kirinyaga Dental measurements (in millimetres) of specimens Laephotis

forest, with ~1900 mm of rainfall per annum), less mesic montane forest (Marsabit National Park) and relatively dry savanna woodlands (e.g. Lolldaiga Hills conservancy FMNH 234558 (other specimens) Table 4. Table Specimen or taxon Holotype kirinyaga Laephotis Laephotis . ‘short-eared’ species of other individuals of the new species and Measurements are of the holotype, range and sample size ( N ). Measurements are presented as the mean ± SD, are taken from Goodman et al. (2017) . malagasyensis , and Lae. matroka robertsi , Lae. Lae. Measurements for the three Malagasy endemics, kirinyaga Laephotis Laephotis capensis Laephotis stanleyi Laephotis robertsi Laephotis matroka Laephotis malagasyensis ~600 mm), hence aridity per se does not seem to be an

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investigation before our hypothesis concerning the geographical range of this species is accepted. The Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 type specimen echolocated at a peak frequency (start and end frequencies) of 44.9 kHz (74.3–41.6 kHz). The mean (± SD) peak frequency for 14 other individuals at the type locality was 43.9 ± 0.91 kHz (73.9 ± 9.43 to 41.8 ± 1.64 kHz).

Pseudoromicia Monadjem, Patterson, Webala & Demos gen. nov. LSID: http://zoobank.org/urn:lsid:zoobank.org:pub: 71737F08-2938-4403-8385-5438B2E5EABE Synonymy Vesperus Peters 1872 (part, not Keyserling & Blasius, 1839). Vesperugo Dobson 1878 (part, not Keyserling & Blasius, 1839). Eptesicus Matschie, 1897 (part, not Rafinesque, 1820). Vespertilio Miller, 1900 (part, not Linnaeus, 1758). Pipistrellus Monard, 1935 (part, not Kaup, 1829). Nycterikaupius (part, not Menu, 1987). Neoromicia Kearney et al., 2002 (part, not Roberts, 1926). Complete synonymic histories for the species placed herein in Pseudoromicia are given in the African Chiroptera report (AfricanBats NPC, 2019).

Type species: Pseudoromicia tenuipinnis (Peters, Figure 8. A, portrait of Laephotis kirinyaga (FMNH 1872). 234558), showing bright brown upper parts and off-white under parts, with bicoloured hairs. The skin around the eye Included species: Pseudoromicia brunnea (Thomas, is blackish in the holotype, but distinctly pinkish in most 1880); Pseudoromicia isabella (Decher, Hutterer & of the paratypes. B, portrait of Pseudoromicia nyanza Monadjem, 2015); Pseudoromicia rendalli (Thomas, (FMNH 215626), showing the distinctive white wings and 1889); Pseudoromicia roseveari (Monadjem et al., under parts of this species. 2013); Pseudoromicia tenuipinnis (Peters, 1872); and two newly described species (see below). important variable in its distribution. However, it has been recorded only at elevations > 1000 m (current Etymology: This feminine name is derived from the records are all between 1160 and 1700 m), and this Greek prefix ψευδο-, false, and the genus Romicia might be an important limit in its geographical Gray, 1838, in turn derived from the Ancient Greek distribution. We also include two specimens (FMNH word ρóμιξα, meaning a ‘kind of javelin or hunting- 233035, 233036) from Murchison Falls National Park, spear’. It also hints at the genus Neoromicia, to which Uganda (1180 m above sea level) in this new species. members of Pseudoromicia were previously assigned. Two specimens from Ethiopia (identified as ‘Neoromicia Members of this new genus resemble and have in the somalica’ by Benda et al., 2016) also group with Lae. past been confused with Neoromicia species. kirinyaga in the phylogeny, as does a specimen from Senegal (Koubínová et al., 2013), suggesting that this Diagnosis: These are small to medium-sized newly described species has a wide distribution north of vespertilionids with a simple muzzle. The tragus is the equator. We recommend, based on its relatively large typically curved anteriorly, with a notch at the base distribution range and habitat preference, that it be listed of the posterior margin. The pelage of the upper and as ‘Least Concern’ in the IUCN red list. However, we did under parts is variably coloured, but in most species not examine the specimens from Ethiopia and Senegal tends to be unicoloured dorsally and bicoloured and therefore recommend a detailed morphological ventrally. In contrast, dorsal pelage is bicoloured in

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 20 A. MONADJEM ET AL.

Afronycteris, Laephotis and Neoromicia. Four of the holotype, and closely resembles it genetically (Fig. 3B) seven species in this genus have translucent white and morphologically (Tables 5–7) and can therefore be Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 wing membranes, whereas membranes are dark considered a paratype. brown or blackish in colour in the remaining three species. The cranium is slightly inflated to relatively Etymology: This species is named in honour of Dr. flattish in lateral profile; in contrast, it is highly Robert M. Kityo, mammalogist, mentor and long- inflated in Afronycteris and moderately inflated in serving curator at the Museum of Zoology, Makerere Neoromicia s.s., whereas it is flattened in Laephotis. University, in recognition of his valuable contributions The outer incisors are usually half the length or less to bats and small mammal research in the region. His of the inner incisors, the latter being weakly bicuspid welcoming nature, curiosity, hospitality and support or unicuspid. The P1 is absent, contrasting with have facilitated numerous and diverse research Afronycteris, in which it is present and relatively agendas over the decades for both national and large. The baculum (~3.0 mm in length) is distinctly international researchers. longer than that of any of the other three genera previously included in Neoromicia, with a robust Diagnosis: This is the largest member of the genus trilobed base and strongly arched shaft leading to a Pseudoromicia, with forearm length of 37 and 38 mm bilobed tip (Fig. 5C). (Table 5) and greatest skull length of 14.70 and 14.99 mm for the two known specimens (Table 6). In Distribution: This genus is widely distributed across comparison, the maximum greatest skull length in sub-Saharan Africa. However, all but one of the species Pse. roseveari (which is the second largest member is associated with equatorial tropical forest and of the genus) is 14.5 mm (Table 6). Pseudoromicia woodland belt. One species, Pse. rendalli, extends far brunnea is smaller in forearm length and in most into savanna habitats, ranging from 13°N to 28°S. craniodental measurements. Therefore, this species is readily diagnosable by size alone. It can easily be distinguished from the white-winged members Systematic relationships: The genera Pseudoromicia of this genus (Pse. rendalli, Pse. isabella and Pse. and Afronycteris are sister to the genera Laephotis and tenuipinnis) by its dark wings. Neoromicia as now understood (see below). Description: External characters: Pseudoromicia kityoi is a large-sized pipistrelle-like bat, similar in Pseudoromicia kityoi Monadjem, Kerbis size to the largest members of the Nycticeinops group, Peterhans, Nalikka, Waswa, Demos & Patterson specifically Nyc. macrocephalus and Nyc. happoldorum, sp. nov. which were both described in the genus Paraphypsugo Kityo’s serotine (Hutterer & Kerbis Peterhans, 2019; Hutterer et al., LSID: http://zoobank.org/urn:lsid:zoobank.org:pub: 2019). Despite its large size, this species is similar in 71737F08-2938-4403-8385-5438B2E5EABE external features to other black-winged members of Holotype: FMNH 223211, field number JCK 7436. This Pseudoromicia. The pelage is medium brown above specimen was collected by Betty Nalikka and Sadic and slightly paler below. The individual hairs are Waswa Babyesiza during a field training exercise with unicoloured on the upper parts and bicoloured on the Julian Kerbis Peterhans. It is an adult male preserved under parts, with the proximal half darker than the in ethanol, with skull extracted and cleaned, and tissue distal half. Like Pse. brunnea and Pse. roseveari, the taken from breast muscle and preserved in dimethyl patagium and uropatagium are both dark in colour. sulfoxide. The ears are short and rounded, and the tragus has a curved outer margin as is typical of the genus Type locality: Mabira Forest Reserve, 0.79 km north- (Monadjem et al., 2013). east of Nagojje Station, Mukono District of the Central Region, Uganda; geographical coordinates: 0.4451°N, Craniodental characters: The skull is robust for a 32.88876°E (Fig. 1). The type specimen was netted on Pseudoromicia, even more so than in Pse. roseveari. 19 October 2012 in cultivated gardens directly adjacent The rostrum has a shallow depression, and the brain (for a photograph of the type locality, see Fig. S2) case is moderately inflated as in other members of to Mabira forest at an elevation of 1130 m above sea the genus. There is no occipital ‘helmet’ as seen in the level. cranium of Lae. capensis (Monadjem et al., 2020b). The sagittal and lambdoidal crests are visible, and the Paratype: One other male (FMNH 223555) was netted zygomatic arches are robust for a pipistrelle-like bat at the same location and on the same night as the (Fig. 9). The dentition in Pse. kityoi is typical of the

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 21 Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 0.29, 3.5–4.4, 3.5–4.4, 4.0 ± 0.29, N = 6 0.87, 4.0–6.0, 4.0–6.0, 4.7 ± 0.87, N = 6 0.94, 4.0–6.8, 4.0–6.8, 5.8 ± 0.94, N = 6 0.99, 4.8–8.3, 4.8–8.3, 6.1 ± 0.99, N = 13 8.3 1.04, 4.8–9.4, 4.8–9.4, 6.0 ± 1.04, N = 15 0.50, 6.0–7.0, 6.0–7.0, 6.5 ± 0.50, N = 10 8.0 7.9 Body mass 1.18, 28.1– 32.0, 28.1– 32.0, 29.4 ± 1.18, N = 6 1.47, 28.0– 31.9, 28.0– 31.9, 30.6 ± 1.47, N = 6 1.30, 33.4–37.0, 33.4–37.0, 34.8 ± 1.30, N = 9 1.24, 29.0– 33.0, 29.0– 33.0, 31.2 ± 1.24, N = 13 31.0 1.06, 32.8–36.7, 32.8–36.7, 34.6 ± 1.06, N = 15 1.54, 32.6–38.0, 32.6–38.0, 36.5 ± 1.54, N = 10 37.0 38.0 Forearm length Forearm 0.91, 12–14, 12–14, 12.7 ± 0.91, N = 7 1.37, 10–14, 10–14, 12.3 ± 1.37, N = 6 1.57, 9–13, 9–13, 11.3 ± 1.57, N = 9 0.64, 11–13, 11–13, 13.0 ± 0.64, N = 13 11 0.83, 11–14, 11–14, 12.4 ± 0.83, N = 15 0.64, 12–14, 12–14, 12.9 ± 0.64, N = 8 10 10 Ear length 0.80, 5–7, 5–7, 6.5 ± 0.80, N = 7 1.18, 5–8, 5–8, 6.9 ± 1.18, N = 6 0.90, 6–9, 6–9, 7.5 ± 0.90, N = 8 0.66, 7–9, 7–9, 7.5 ± 0.66, N = 13 7 0.97, 7–10, 7–10, 7.79 ± 0.97, N = 14 1.10, 8–11, 8–11, 9.9 ± 1.10, N = 10 9 10 Hindfoot length 0.84, 28–30, 28–30, 29.2 ± 0.84, N = 5 4.23, 24–36, 24–36, 29.3 ± 4.23, N = 6 3.15, 32–42, 32–42, 37.0 ± 3.15, N = 9 2.29, 30–38, 30–38, 33.3 ± 2.29, N = 13 30 1.55, 33–38, 33–38, 35.9 ± 1.55, N = 15 2.65, 34–44, 34–44, 39.7 ± 2.65, N = 9 36 34 Tail length Tail 2.17, 72–77, 72–77, 74.2 ± 2.17, N = 5 2.73, 74–82, 74–82, 78.8 ± 2.73, N = 15 3.92, 82–95, 82–95, 88.5 ± 3.92, N = 15 2.73, 79–89, 79–89, 83.5 ± 2.73, N = 13 83 3.14, 80–89, 80–89, 84.8 ± 3.14, N = 15 2.95, 83–93, 83–93, 87.3 ± 2.95, N = 10 89 88 Total length Total

Pseudoromicia Pseudoromicia Uganda and Reserve, kityoi from Mabira Forest External measurements (in millimetres) and mass grams) of Pseudoromicia

(West Africa) (West (other specimens) Holotype FMNH 215626 Paratype FMNH 223555 Paratype Holotype FMNH 223211 Pseudoromicia tenuipinnis s.s. Pseudoromicia isabella Pseudoromicia rendalli nyanza Pseudoromicia nyanza Pseudoromicia Pseudoromicia brunnea Pseudoromicia roseveari Pseudoromicia kityoi Pseudoromicia Pseudoromicia kityoi Pseudoromicia from Kisumu, Kenya from Kisumu, nyanza Table 5. Table Specimen or taxon The . Pseudoromicia other individuals of the two new species and Measurements are of the holotypes, range and sample size ( N ). Measurements are presented as the mean ± SD, the four below are white winged (see main text for more details). line are dark winged, three species listed above the horizontal black

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 22 A. MONADJEM ET AL. Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 8.97 MAND 11.25 10.76 0.30, 10.40 ± 0.30, 9.80–10.70, N = 9 0.32, 10.06 ± 0.32, 9.40–10.66, N = 14 0.19, 9.06 ± 0.19, 8.80–9.53, N = 18 0.33, 9.95 ± 0.33, 9.40–10.50, N = 12 0.20, 9.64 ± 0.20, 9.32–9.87, N = 5 0.38, 8.77 ± 0.38, 8.20–9.33, N = 10 GSH 5.46 5.38 0.12, 5.21 ± 0.12, 5.10–5.40, N = 7 0.25, 5.14 ± 0.25, 4.80–5.50, N = 8 5.05 0.17, 4.81 ± 0.17, 4.54–5.14, N = 17 0.27, 5.05 ± 0.27, 4.70–5.57, N = 8 0.09, 4.97 ± 0.09, 4.88–5.06, N = 3 0.34, 4.68 ± 0.34, 4.40–5.41, N = 7 GBW 7.67 7.51 0.35, 7.31 ± 0.35, 6.87–7.40, N = 10 0.26, 7.08 ± 0.26, 6.65–7.69, N = 15 6.90 0.20, 6.95 ± 0.20, 6.73–7.45, N = 17 0.31, 7.25 ± 0.31, 6.80–7.80, N = 12 0.16, 6.77 ± 0.16, 6.53–6.92, N = 5 0.13, 6.51 ± 0.13, 6.40–6.81, N = 10 MAST 8.25 8.19 0.26, 7.84 ± 0.26, 7.50–8.40, N = 10 0.24, 7.50 ± 0.24, 7.10–7.93, N = 15 7.31 0.21, 7.32 ± 0.21, 6.99–7.76, N = 17 0.32, 7.76 ± 0.32, 7.18–8.33, N = 12 0.19, 7.27 ± 0.19, 6.96–7.48, N = 5 0.14, 6.88 ± 0.14, 6.75–7.22, N = 10 POB 3.99 3.91 0.19, 3.80 ± 0.19, 3.50–4.09, N = 10 0.23, 3.92 ± 0.23, 3.60–4.45, N = 15 3.69 0.23, 4.00 ± 0.23, 3.69–4.70, N = 17 0.13, 3.98 ± 0.13, 3.80–4.17, N = 12 0.15, 3.68 ± 0.15, 3.50–3.89, N = 5 0.20, 3.79 ± 0.20, 3.47–4.25, N = 10 ZYGO 9.63 9.54 0.50, 8.85 ± 0.50, 8.00–9.50, N = 9 0.49, 8.68 ± 0.49, 7.70–9.61, N = 14 8.12 0.30, 8.06 ± 0.30, 7.54–8.69, N = 17 0.42, 9.05 ± 0.42, 8.40–9.90, N = 10 0.43, 8.28 ± 0.43, 7.58–8.65, N = 5 0.50, 7.48 ± 0.50, 6.80–8.19, N = 9 GSKL 14.99 14.70 0.45, 14.13 ± 0.45, 13.40–14.50, N = 10 0.35, 13.73 ± 0.35, 13.10–14.24, N = 15 13.20 0.25, 12.96 ± 0.25, 12.48–13.43, N = 17 0.48, 13.57 ± 0.48, 12.98–14.70, N = 11 0.15, 13.12 ± 0.15, 12.99–13.34, N = 5 0.28, 12.43 ± 0.28, 12.00–12.78, N = 10 from nyanza Uganda and Pseudoromicia Reserve, kityoi from Mabira Forest Cranial measurements (in millimetres) of specimens Pseudoromicia

(West Africa) (West Holotype FMNH 223211 Paratype FMNH 223555 Paratype Holotype FMNH 215626 (other specimens) s.s. Kisumu, Kenya Kisumu, Table 6. Table Specimen or taxon The . Pseudoromicia other individuals of the two new species and Measurements are of the holotypes, range and sample size ( N ). Measurements are presented as the mean ± SD, the four below are white winged (see main text for more details). line are dark winged, three species listed above the horizontal black Pseudoromicia kityoi Pseudoromicia Pseudoromicia kityoi Pseudoromicia Pseudoromicia roseveari Pseudoromicia brunnea nyanza Pseudoromicia nyanza Pseudoromicia Pseudoromicia rendalli Pseudoromicia isabella Pseudoromicia tenuipinnis

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 23

Table 7. Dental measurements (in millimetres) of specimens of Pseudoromicia kityoi from Mabira Forest Reserve, Uganda and Pseudoromicia nyanza from Kisumu, Kenya Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020

3 3 3 Specimen or taxon C–M C–C M –M c–m3

Pseudoromicia kityoi 5.20 4.74 6.10 5.63 Holotype FMNH 223211 Pseudoromicia kityoi 5.12 4.61 6.09 5.59 Paratype FMNH 223555 Pseudoromicia roseveari 5.03 ± 0.20, 4.34 ± 0.25, 6.00 ± 0.25, 5.25 ± 0.32, 4.80–5.30, N = 10 3.80–4.70, N = 10 5.50–6.30, N = 10 5.00–5.97, N = 9 Pseudoromicia brunnea 4.86 ± 0.13, 4.22 ± 0.24, 5.86 ± 0.26, 5.30 ± 0.33, 4.60–5.07, N = 15 3.63–4.63, N = 15 5.50–6.55, N = 15 4.80–5.86, N = 15 Pseudoromicia nyanza 4.31 4.19 5.26 4.62 Holotype FMNH 215626 Pseudoromicia nyanza 4.27 ± 0.10, 4.07 ± 0.11, 5.24 ± 0.17, 4.62 ± 0.11, (other specimens) 4.04–4.46, N = 18 3.89–4.29, N = 18 4.93–5.49, N = 18 4.42–4.80, N = 18 Pseudoromicia rendalli 4.68 ± 0.18, 4.38 ± 0.25, 5.65 ± 0.32, 5.07 ± 0.22, 4.44–5.10, N = 12 4.00–4.80, N = 12 5.10–6.22, N = 10 4.60–5.49, N = 12 Pseudoromicia isabella 4.49 ± 0.05, 4.40 ± 0.18, 5.51 ± 0.11, 5.04 ± 0.19, 4.43–4.56, N = 5 4.15–4.59, N = 5 5.33–5.61, N = 5 4.86–5.35, N = 5 Pseudoromicia tenuipinnis 4.23 ± 0.14, 3.81 ± 0.29, 5.03 ± 0.18, 4.63 ± 0.23, s.s. (West Africa) 3.90–4.35, N = 10 3.30–4.14, N = 9 4.70–5.29, N = 9 4.40–5.24, N = 10

Measurements are presented as the mean ± SD, range and sample size (N). Measurements are of the holotypes, other individuals of the two new species and other species of Pseudoromicia. The three species listed above the horizontal black line are dark winged, the four below are white winged (see main text for more details). genus, with I 2/3, C 1/1, P 1/3, M2/3. In the upper tooth with rapidly disappearing forest habitat, this species is row, I1 is unicuspid and I2 is tiny, extending slightly probably of conservation concern. beyond the cingulum of I1. The P1 is absent, putting Its closest known relative is Pse. roseveari, recently 2 C in contact with P . The m3 is myotodont sensu Van described from Mount Nimba and with a limited Cakenberghe & Happold (2013). distribution in the borderland zone between Liberia and Guinea (Monadjem et al., 2013; Decher et al., 2015; Mamba et al., in press), some 4700 km to the west. Biology: Owing to the paucity of specimens, almost Whether either species occurs in the vast tropical nothing can be said about the biology of this species. rainforests between these two sites is unknown and The only two known specimens were captured within deserves investigation. 200 m from the edge of Mabira Forest in a domestic garden (Supporting Information, Fig. S2). However, considering that most members of this genus are Pseudoromicia nyanza Monadjem, Patterson, restricted to tropical rainforest habitats, and that the Webala & Demos sp. nov. two known specimens of this species were captured in Nyanza serotine a remnant patch of rainforest, its global distribution LSID: http://zoobank.org/urn:lsid:zoobank.org:pub: might be both fragmented and limited in extent. 71737F08-2938-4403-8385-5438B2E5EABE Urgent surveys are required to assess the status of Neoromicia tenuipinnis Patterson & Webala (2012). this species at Mabira Forest Reserve, which has been Neoromicia tenuipinnis Musila et al. (2019). steadily losing habitat to agriculture over the past few Neoromicia tenuipinnis Rydell et al. (2020). decades (Boffa et al., 2008). We suggest that this species might be present in other Congo Basin forest patches Holotype: FMNH 215626, field number BDP 5719. in Uganda (e.g. Semliki, Kibale, Kashyoha-Kitomi) and This specimen was collected on 8 January 2012 by Kenya (Kakamega), although extensive surveys at Bruce D. Patterson, Paul W. Webala and Carl W. Dick. Kakamega forest have failed to locate this species there It is an adult male, formalin-fixed and preserved in (Webala et al., 2019). Owing to the limited information ethanol. Its skull has been extracted and cleaned, its available on this species, we recommend that it be given glans penis removed and the baculum stained and the IUCN conservation status of ‘Data Deficient’, but extracted. Muscle tissue was also preserved in liquid we note that because of its presumed close association nitrogen at the time of capture.

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 24 A. MONADJEM ET AL.

other Pseudoromicia species (Fig. 3B). Furthermore, it is readily distinguished from the dark-winged members Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 of this genus (Pse. roseveari, Pse. brunnea and Pse. kityoi) by its white wings. It can be distinguished from Pse. rendalli by its smaller size (mostly non-overlapping forearm length and craniodental measurements (Tables 5–7) and weakly bicuspid I1 (unicuspid in Pse. rendalli). It is significantly larger than Pse. tenuipinnis, with hardly any overlapping external and craniodental measurements (Tables 5–7); furthermore, its dorsal fur is medium brown and bicoloured (dark brown and unicoloured in Pse. tenuipinnis). It is most like Pse. isabella in size and external appearance, but that species has rusty tips to the fur on its upper parts, whereas Pse. nyanza has white-tipped hairs. The taxon Eptesicus ater J. A. Allen, 1917, which was described from north-eastern Democratic Republic of the Congo, is currently considered a synonym of Pse. tenuipinnis (Simmons, 2005) and is far smaller than Pse. nyanza, with a reported total length of 68 mm. Furthermore, Pse. tenuipinnis has ‘brownish black’ fur on its back (Allen et al., 1917), contrasting with the light-tipped fur of Pse. nyanza. Figure 9. Plate showing the cranium and mandible of Pseudoromicia kityoi (FMNH 223211). Scale bar = 5 mm. Description: External characters: Pseudoromicia nyanza is a medium-sized pipistrelle-like bat with Type locality: Kisumu Impala Sanctuary, State white patagial and uropatagial membranes (Fig. 8B). Lodge Campsite, Kisumu County (formerly Nyanza The dorsal pelage is medium brown with white-tipped province), Kenya, at an elevation of 1130 m above sea hairs over most of the back. The ventral hairs are level; geographical coordinates: 0.10961°S, 34.74593°E pure white with a dark base. The ears are short and (Fig. 1). The sanctuary borders both Lake Victoria and rounded, and the tragus is broad and truncated, as in Kenya’s fifth largest city, Kisumu, and is only 0.34 km2 Pse. tenuipinnis (Monadjem et al., 2013). in area. Vegetation consisted of open parkland, short- statured trees and shrubs. Craniodental characters: The skull is relatively gracile, as in Pse. tenuipinnis and Pse. isabella. In Paratypes: Four other individuals (FMNH 215625, lateral profile, the cranium slopes gently up from FMNH 215627, FMNH 215628 and FMNH 215629), the rostrum to the top of the braincase. There is no all females, were collected at the same location and on occipital ‘helmet’, and the sagittal and lambdoidal the same night as the holotype and closely resemble it crests are absent. The zygomatic arches are fragile, genetically (Fig. 3B) and morphologically (Tables 5–7), as in Pse. tenuipinnis and Pse. isabella (Fig. 10). The qualifying them as paratypes. dentition in Pse. nyanza is typical of the genus, with I 2/3, C 1/1, P 1/3, M 2/3. In the upper tooth row, I1 Etymology: This species is named after the region is weakly but distinctly bicuspid and I2 is moderate where it was found, Nyanza, which derives from the in size, slightly more than half the length of I1. The 1 2 Bantu word for ‘large body of water’. Covering nearly P is absent, putting C in contact with P . The m3 is 60 000 km2, Lake Victoria surely qualifies. The name is myotodont sensu Van Cakenberghe & Happold (2013). used as a noun in apposition. Biology: Judging by how frequently this species Diagnosis: This is a medium-sized member of the is captured, it is common west of the Rift Valley in genus Pseudoromicia, with a mean forearm length Kenya (B. D. Patterson & P. W. Webala, personal of 31.2 mm (Table 5) and greatest skull length of observation). It seems to prefer forest-edge habitats 12.96 mm (Table 6). It is genetically distinct from all and avoids the forest interior (Rydell et al., 2020,

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 25

Afronycteris Monadjem, Patterson & Demos gen. nov. Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 LSID: http://zoobank.org/urn:lsid:zoobank.org:pub: 71737F08-2938-4403-8385-5438B2E5EABE

Synonymy Vespertilio Peters, 1852 (part, not Linnaeus, 1758). Hypsugo Kolenati, 1860 (part, not Kolenati, 1856). Vesperugo Dobson, 1875 (part, not Keyserling & Blasius, 1839). Pipistrellus Miller, 1900 (part, not Kaup, 1829). Myotis Matschie, 1907 (part, not Kaup, 1829). Neoromicia Shortridge, 1934 (part, not Roberts, 1926). Eptesicops Roberts, 1951 (part, not Roberts, 1926). Complete synonymic histories for the species placed herein in Afronycteris are given in the African Chiroptera report (AfricanBats NPC, 2019).

Type species: Afronycteris nana (Peters, 1852).

Included species: Afronycteris helios (Heller, 1912). Figure 10. Plate showing the cranium and mandible of Pseudoromicia nyanza (FMNH 215626). Scale bar = 5 mm. Etymology: From the Greek word νυχτερίδα, bat, and the prefix Afro- referring to the African continent, as Neo. tenuipinnis). However, its distribution referring to the wide distribution of the type species beyond western Kenya is not known. It seems to be A. nana. This species ranges, without obvious breaks in associated with the high plateau of western Kenya, distribution, from Senegal in the west, east to Ethiopia which extends into eastern Uganda; presumably, it and south to South Africa, being absent only from also occurs there. Thorn & Kerbis Peterhans (2009) the more arid desert and semi-desert environments recorded ‘Pipistrellus tenuipinnis’ as occurring widely associated with the Sahara, Sahel and Chalbi Desert in Uganda. The cranial measurements of specimens in the north and the Namib and Kalahari deserts in from Budongo, Entebbe and Sango Bay (at elevations the south-west (Happold, 2013a). similar to those we report from Kenya) all fall neatly within the range of Pse. nyanza and are generally Diagnosis: Small-sized vespertilionids with the simple larger than those for Pse. tenuipinnis. It would be muzzle characteristic of this family. The cranium in instructive to re-examine these specimens (in the lateral view is distinctly inflated, more so than any collections of the BMNH and LACM) to confirm their other member of the tribe Vespertilionini. The tragus identities and help to determine the western limits is characteristically hatchet shaped, with the posterior of the distribution of Pse. nyanza. However, records margin having an abrupt angle and lacking a notch from the eastern Democratic Republic of the Congo at its base, as illustrated by Van Cakenberghe & apparently refer to true Pse. tenuipinnis, owing Happold (2013). The tragi of Laephotis, Neoromicia to their small size, with total length ‘about 72 mm’ and Pseudoromicia all have a notch at the base of the (Allen et al., 1917). We speculate that, despite the posterior margin. The pelage of the upper and under limited geographical range of Pse. nyanza (even if parts is bicoloured, with the basal portion of each hair Uganda is included), this species is currently not darker than the terminal portion. There is a distinct threatened because it survives in human-altered thumbpad at the base of the thumb, thought to be habitats, and therefore we recommend the IUCN useful in climbing on smooth leaves. The outer incisor conservation status of ‘Least Concern’. The type I2 is well developed, reaching almost the same length specimen echolocated at a peak frequency (start and as the I1, the latter being slightly bicuspid or unicuspid; end frequencies) of 40.4 kHz (56.4–39.3 kHz). The in Laephotis, Neoromicia and Pseudoromicia, I2 is mean (± SD) peak frequency for 16 individuals at typically half the length of I1 or shorter. The P1 is present the type locality was 40.4 ± 0.84 kHz (55.1 ± 7.91 to and relatively large, whereas this tooth is absent in 39.5 ± 0.68 kHz). Laephotis, Neoromicia (except Neo. bemainty and Neo.

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 26 A. MONADJEM ET AL. anchietae) and Pseudoromicia. The baculum (~2.0 mm species of Laephotis in a single undifferentiated in length) is shorter than in Pseudoromicia and similar genus; (2) to treat Laephotis, Neoromicia and the Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 in length to that of Laephotis and Neoromicia. It has a newly named groups Afronycteris and Pseudoromicia distinctly and deeply bilobed base and a gently curved as subgenera; or (3) to treat all four clades as distinct shaft leading to a spatulate tip (Fig. 5D). genera. Assigning ranks to clades is always a subjective task. Given that zoological nomenclature serves both Distribution: This genus is endemic to sub-Saharan information storage and retrieval functions (Mayr, Africa, probably occurring in suitable habitats across 1969), it is important that rank assignments be its wide range. It occurs throughout the Upper more or less equivalent among comparable groups Guinea rainforest zone, extending northward into and that systematists strive to conserve the stability Sudanian savanna, possibly extending into the Sahel of binomial nomenclature insofar as possible. along major rivers and wetlands (Happold, 2013a). It Morphological discontinuities, genetic distances and occurs throughout mesic portions of Central and East chronological ages have all been used in determination Africa, but records are sparser in the Horn of Africa of group ranks. Here, we note that each of the genera (Lanza et al., 2015). It is widespread in the wetter we recognize is distinguished by trenchant genetic parts of southern Africa, avoiding the dry south- distances (Table 1) and morphological discontinuities western region of South Africa, much of Botswana (e.g. Fig. 5) that are comparable to those among other and Namibia (Monadjem et al., 2010). recognized genera of Vespertilionini. Furthermore, the nomenclatural history and phylogenetic relationships Systematic relationships: Afronycteris is sister to of the group make it impossible to conserve traditional Pseudoromicia, but the two genera can be distinguished binomial usage. easily by external characteristics, cranial features and There is renewed attention being paid to the utility the shape of the baculum (see ‘Diagnosis’ above for of the subgenus category as a means to incorporate details). phylogenetic information without disrupting binomial usage (Voss et al., 2014; Teta, 2018). Yet in our case, both options 1 (one undifferentiated genus) and 2 (the use of subgenera) would only complicate matters DISCUSSION for other biologists using scientific nomenclature. We review nearly all the sub-Saharan pipistrelle-like Laephotis Thomas, 1901 has priority over Neoromicia bats in the tribes Vespertilionini and Pipistrellini. Roberts, 1926 or other later proposed names. Within this region, the tribe Pipistrellini is Therefore, option 1 entails recognition of all these represented by the sister genera Pipistrellus and species in the genus Laephotis and would thus change Scotoecus, with V. rueppellii (formerly considered as a the generic assignment of all 17 species currently member of Pipistrellus) sister to these two. Vansonia recognized in Neoromicia. In turn, option 2 would must be considered a valid genus (Koubínová cause the same disruption plus levy the additional et al., 2013; Moratelli & Burgin, 2019), because nomenclatural burden of subgenera. Our proposal Scotoecus, Nyctalus Bowdich, 1825 and Glischropus (option 3) conserves the usage of both Laephotis and Dobson, 1875 would otherwise render Pipistrellus Neoromicia insofar as possible, expanding previous paraphyletic. None of these three genera is endemic concepts of the former to include short-eared forms to sub-Saharan Africa (or even to Africa; Moratelli and restricting application of the latter to forms more & Burgin, 2019). Based on the cytochrome b gene, closely related to Neo. zuluensis. It underscores the Mimetillus Thomas, 1904 does not fall into either discovery of a distinctive, largely white-winged clade of these two tribes, but is sister to them. It was that is mainly restricted to tropical forests with the placed in Vespertilionini by Simmons (2005), but new name Pseudoromicia and highlights the phyletic this is the first time that it has been included in a remoteness of the ubiquitous banana bat by placing it comprehensive molecular phylogeny of these two (or them) in the genus Afronycteris. tribes. Additional genetic data are needed to place In our new conception, the genus Laephotis includes and classify Mimetillus securely. the long-eared species traditionally recognized by that Generic delimitation within the tribe Vespertilionini name (Lae. angolensis, Lae. botswanae, Lae. namibensis is both subtle and complicated, as amply illustrated by and Lae. wintoni) and what was previously called its tortuous synonymic history (Table 8). In resolving Neo. capensis and allied species (all short-eared by the nomenclature of Neoromicia as traditionally comparison). Based on our phylogeny, the long-eared conceived, we were faced with three options: (1) to species of Laephotis are sister to a group comprising combine all members of this group plus the four known (Lae. capensis + Lae. matroka) + Lae. kirinyaga. The

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 27

Table 8. Recent changes to the taxonomy of African and Malagasy Vespertilionini and Pipistrellini Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020

Simmons (2005)* Simmons & Cirranello (2020) This study

Neoromicia nanus Laephotis nanus Afronycteris nana Pipistrellus helios Laephotis helios (Afronycteris helios)† Hypsugo ariel Hypsugo ariel Hypsugo ariel Hypsugo musciculus Hypsugo musciculus (Hypsugo musciculus) Laephotis angolensis Laephotis angolensis (Laephotis angolensis) Laephotis botswanae Laephotis botswanae Laephotis botswanae Neoromicia capensis Laephotis capensis Laephotis capensis – – Laephotis kirinyaga Neoromicia somalicus malagasyiensis Laephotis malagasyiensis Laephotis malagasyiensis Neoromicia capensis Laephotis matroka Laephotis matroka Laephotis namibensis Laephotis namibensis Laephotis namibensis – Laephotis robertsi Laephotis robertsi – Laephotis stanleyi Laephotis stanleyi Laephotis wintoni Laephotis wintoni Laephotis wintoni – – [Laephotis cf. wintoni] Hypsugo anchietae Laephotis anchietae (Neoromicia anchietae)‡ – Hypsugo bemainty Neoromicia bemainty Neoromicia guineensis Laephotis guineensis (Neoromicia guineensis)§ Neoromicia somalica Laephotis somalicus Neoromicia somalica – – [Neoromicia cf. somalica] Neoromicia zuluensis Laephotis zuluensis Neoromicia zuluensis Hypsugo crassulus bellieri Parahypsugo bellieri Nycticeinops bellieri Hypsugo crassulus crassulus Parahypsugo crassulus Nycticeinops crassulus Hypsugo eisentrauti Parahypsugo eisentrauti Nycticeinops eisentrauti Neoromicia capensis grandidieri Pipistrellus grandidieri Nycticeinops grandidieri – Parahypsugo happoldorum Nycticeinops happoldorum – Parahypsugo macrocephalus (Nycticeinops macrocephalus)¶ Nycticeinops schlieffeni Nycticeinops schlieffeni Nycticeinops schlieffeni – – [Nycticeinops cf. schlieffeni] Neoromicia brunnea Laephotis brunneus Pseudoromicia brunnea – Laephotis isabella Pseudoromicia isabella – – Pseudoromicia kityoi – – Pseudoromicia nyanza Neoromicia rendalli Laephotis rendalli Pseudoromicia rendalli – Laephotis roseveari Pseudoromicia roseveari Neoromicica tenuipinnis Laephotis tenuipinnis Pseudoromicia tenuipinnis Pipistrellus areo Pipistrellus areo (Pipistrellus areo) Pipistrellus hesperidus Pipistrellus hesperidus Pipistrellus hesperidus – – [Pipistrellus cf. hesperidus] Pipistrellus inexspectatus Pipistrellus inexspectatus (Pipistrellus inexspectatus) Pipistrellus nanulus Pipistrellus nanulus Pipistrellus nanulus Pipistrellus permixtus Pipistrellus permixtus (Pipistrellus permixtus) Pipistrellus raceyi Pipistrellus raceyi Pipistrellus raceyi Pipistrellus rusticus Pipistrellus rusticus (Pipistrellus rusticus) – – Pipistrellus simandouensis Scotoecus albigula Scotoecus albigula (Scotoecus albigula) Scotoecus albofuscus Scotoecus albofuscus (Scotoecus albofuscus) Scotoecus hindei Scotoecus hindei Scotoecus hindei Scotoecus hirundo Scotoecus hirundo Scotoecus hirundo Pipistrellus rueppellii Pipistrellus rueppellii Vansonia rueppellii

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Table 8. Continued Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 Simmons (2005)* Simmons & Cirranello (2020) This study

– – [Vansonia cf. rueppellii] Mimetillus moloneyi Mimetillus moloneyi#

Taxa listed in parentheses need genetic confirmation of their generic allocation; those listed in square brackets require careful delimitation and taxonomic description. *Two additional species listed were Neoromicia melckorum, now regarded as a synonym of Laephotis capensis (Goodman et al., 2017), and Neoromicia flavescens, now considered a nomen dubium (see Thorn et al., 2007). †See Happold & Van Cakenberghe (2013) for problems with this name. ‡Allocation based on its close genetic relationship with Neo. bemainty. §Allocation based on its bacular morphology. ¶Allocation based on its morphological similarity to Nyc. happoldorum. #Shown by our genetic analysis to fall outside Vespertilionini and Pipistrellini. clade of Lae. robertsi + Lae. malagasyensis is sister to Guinea zone of West Africa, where up to four species all these above-mentioned species, and the recently can co-occur in the same patch of forest (Monadjem described Lae. stanleyi is sister to this broader group. et al., 2016). One species (Pse. rendalli) is associated The close relationship between the long-eared Laephotis with wetlands outside forested habitats and ranges and some species formerly recognized as Neoromicia widely across the continent with little geographical has long been noted (Hoofer & Van den Bussche, 2003) structuring. Interestingly, the white-winged form and discussed (e.g. Roehrs et al. 2010; Koubínová et al. present in western Kenya and previously identified 2013). Furthermore, this is not a unique instance where as Neoromicia tenuipinnis Peters, 1872 (Musila et al., a distinctive long-eared vesper taxon has been shown 2019) is not closely related to Pse. tenuipinnis s.s. (as to be deeply embedded in an otherwise short-eared defined here). Additionally, a specimen collected from group. The Neotropical genus-group Histiotus Gervais, Minziro Forest, north-west Tanzania and previously 1856 renders even New World members of the genus identified as Neo. tenuipinnis (Stanley & Foley, 2008) Eptesicus Rafinesque, 1820 paraphyletic (Amador does not group with our newly described species of et al., 2018). Interestingly, bacular morphology (see western Kenya, Pse. nyanza. In fact, it is highly distinct Fig. 5) supports our revised definition of this genus, genetically and might belong to an undescribed because both long-eared and short-eared members species. Two of the three known dark-winged species have similar bacular morphologies (Hill & Harrison, occur in West Africa, with Pse. brunnea ranging into 1987). Skull morphometrics also distinguish them as the Lower Guinea forests of Cameroon (Fahr, 2013). a group (Fig. 4). This suggests that the lengthening The third species is described here as Pse. kityoi, of ear pinnae in vespertilionid bats might occur which is currently known from only a small forest in rapidly, possibly as an adaptive response (e.g. for central Uganda; additional surveys might show it to gleaning prey off the ground) and might not be a good range more widely in the region. character in defining generic limits. All members of The widespread and relatively abundant species the more expansive genus Laephotis dictated by our previously identified as Neo. nana (and the poorly analyses are endemic to sub-Saharan Africa (including understood Neo. helios) is not similar to any of Madagascar). these other groups. We demonstrate that it is highly Neoromicia as we refine its application here distinct genetically and is sister to Pseudoromicia. comprises the type species of the genus, Neo. zuluensis, Furthermore, it has a suite of unique characters not and its sister, Neo. somalica. In addition, we include shared with members of the genus Pseudoromicia, Neo. bemainty and Neo. anchietae in this genus, a such as an inflated cranium, disc pads at the base of its relationship also noted previously (Goodman et al., thumb, the presence of a small upper premolar, a large 2015, 2017). All these species are relatively small- and long outer upper incisor and a uniquely structured bodied forms with a similar three-pronged (cross- baculum. It belongs in its own genus, which we have shaped) tip to the baculum (Hill & Harrison, 1987), named Afronycteris. unique to this genus. In addition to Afronycteris, Laephotis, Neoromicia and The newly described genus Pseudoromicia comprises Pseudoromicia, the following taxa are also documented a group mostly associated with African tropical forests. in the sub-Saharan region: Afropipistrellus, Hypsugo Many species possess highly distinctive white wings, and Nycticeinops. Hypsugo is represented in the sub- which, judged from our phylogeny, appear ancestral for Saharan region only by Hypsugo ariel (Thomas, 1904), this genus. It has the highest diversity in the Upper based on a single record in Sudan (Koopman, 1975),

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 A REVISION OF PIPISTRELLE-LIKE BATS 29 and is unlikely to be widespread beyond the arid species (Monadjem et al., 2010, 2020b), suggesting Sahel. In contrast, Afropipistrellus comprises a purely that the former also belongs to Neoromicia. Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 African lineage mostly associated with rainforest; Pipistrellus aero is not represented in our phylogeny, one species (Afr. grandidieri) is associated with moist because we did not sample at the type locality and woodlands (Monadjem et al., 2020a). The rainforest no DNA sequences of this species are available members of this group were recently described in the on GenBank. We could not include topotypical new genus Parahypsugo (Hutterer et al., 2019), but Parahypsugo crassulus, but we suspect from its close genetic material now available for Afr. grandidieri, type morphological similarity to Nyc. bellieri (De Vree, species of Afropipistrellus (Thorn et al., 2007), shows 1972) that it will eventually be shown to belong to that it clearly falls into this group, and Afropipistrellus Nycticeinops. Furthermore, we included a specimen has priority over Parahypsugo. Our alignment also from Tanzania in our phylogeny that we tentatively included a 762 bp Cytb sequence from an individual identified on craniodental grounds as Parahypsugo identified as Par. eisentrauti. This individual was crassulus (Fig. 3B). This specimen clearly groups not recovered with Afropipistrellus, which grouped with other members of Afropipistrellus and further instead with Nycticeinops. As an expression of our supports the contention that Par. crassulus belongs taxonomic conservatism, we regard Parahypsugo as in the genus Nycticeinops. a junior synonym of Afropipistrellus and synonymize Specimens from Yemen identified as Neo. the latter with Nycticeinops, provisionally including guineensis (Benda et al., 2011; Juste et al., 2013) are the anomalous sequence reported for Par. eisentrauti. clearly sister to Neo. somalica, but Neo. guineensis This transforms Nycticeinops, which is traditionally has yet to be sequenced at its type locality (Guinea regarded as monospecific, into a genus containing Bissau), and the Yemeni specimens might represent at least seven species. We note that the intrageneric a different species. We are not sure whether distance within Nycticeinops is relatively large our samples include Pip. helios. Judging by the compared with other genera within the Pipistrellini apparent deep divisions within the clade (with an and Vespertilionini but is still lower than the intraspecific cytochrome b divergence of 3.2%; see intergeneric distances (see Table 1). In this genus, the Supporting Information Table S4) that we have substantial, geographically structured differentiation named Afr. nana, more than one species might be of Nyc. schlieffeni into eastern and western clades involved. The relationship between Afr. helios and (also noted by Koubínová et al., 2013) and of Nyc. Afr. nana requires further study. We were unable grandidieri into eastern and southern African clades to include Pipistrellus inexspectatus Aellen, 1959 in deserves further study. our tree. However, a recent study based on the COI Including the three new species described in this gene suggested that this species might not belong in paper, there are now 46 species of Vespertilionini and Pipistrellus at all, although it was unclear whether Pipistrellini with valid names recorded from sub- true Pip. inexspectatus had been sampled (Monadjem Saharan Africa (Table 8), five of which are endemic to et al., 2020a). Finally, we were also lacking genetic Madagascar (Goodman, 2011; Goodman et al., 2012, material from either Hypsugo musciculus Thomas, 2015). Of the remaining 41 species, 15 (or more than 1913 or Pipistrellus permixtus Aellen, 1957 (the one-third) of them have been recorded from Kenya latter known only from the holotype from Dar-es- and 20 (49%) species from East Africa, demonstrating Salam, Tanzania), meaning that we cannot speculate the importance of this region for pipistrelle-like bat about their generic relationships. It is worth noting diversity. This corroborates previous studies showing that the origins of Pip. permixtus have been disputed Kenya to be a hub of genetic diversity for other bat because it is morphologically more similar to that groups, such as the species-rich genera Hipposideros of Palaearctic and Oriental members of the genus, Gray, 1831, Miniopterus Bonaparte, 1837, Rhinolophus such as Pip. pipistrellus (Schreber, 1774) and Pip. Lacépède, 1799 and Scotophilus Leach, 1821 (Demos nathusii (Keyserling & Blasius, 1839), than to any et al., 2018, 2019a, b, 2020; Patterson et al., 2020) and African species (Aellen, 1957; Happold, 2013b). further emphasizes the need for continued taxonomic In conclusion, based on extensive genetic sampling surveys in the region. and morphological investigation, we have taken We were not able to gather genetic material for important steps towards resolving the systematic all sub-Saharan species of the tribes Pipistrellini relationships of a poorly understood group of and Vespertilionini and therefore leave a number pipistrelle-like bats in the tribes Vespertilionini and of taxonomic issues outstanding (Table 8). We did Pipistrellini in sub-Saharan Africa. Furthermore, not include Vesperugo anchietae Seabra, 1900 in we have addressed pending taxonomic issues by our phylogeny, but we did include Neo. bemainty describing two new genera and three new species (Goodman et al., 2015), which is a closely related within the Vespertilionini.

© 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2020, XX, 1–33 30 A. MONADJEM ET AL.

ACKNOWLEDGEMENTS with a review of related Vespertilioninae from the island. Acta Chiropterologica 8: 299–324. Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa087/5903787 by University of New England user on 10 September 2020 We thank Carl Dick, Beryl Makori, Ruth Makena, Benda P, Al-Jumaily MM, Reiter A, Nasher AK. 2011. Bernard Agwanda, David Wechuli, Richard Yego, Noteworthy records of bats from Yemen with description Michael Bartonjo and Aziza Zuhura for help in of a new species from Socotra. Hystrix, Italian Journal of obtaining specimens in the field. We are grateful for the Mammalogy 22: 23–56. efforts of curators and collection managers in all the Benda P, Reiter A, Uhrin M, Varadinová Z. 2016. A new institutions cited in the Supporting Information (Table species of pipistrelle bat (Chiroptera: Vespertilionidae) from S1) for maintaining the museum voucher specimens southern Arabia. 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SUPPORTING INFORMATION Additional Supporting Information may be found in the online version of this article at the publisher’s web-site: Table S1. Bat specimens referred to in this study. We present the museum where known. The ‘y’ refers to a specimen in which cytochrome b (Cytb) was sequenced in this study and/or a specimen that was measured in this study (‘Morphology’). ‘Redundant’ refers to specimens that were sequenced in this study but not included in the phylogenetic tree (Fig. 3) because they would have cluttered the tree unnecessarily. Specimens, which were sequenced by other studies and for which cytochrome b sequences were obtained from GenBank are provided with the GenBank accession numbers. Where known, the locality and latitude/longitude are presented. Museum abbreviations are as follows: EMNH, Eswatini National Museum of Natural History, Kwaluseni; EOWL, E. O. Wilson Biodiversity Laboratory, Chitengo; FMNH, The Field Museum of Natural History, Chicago; HNHM, Hungarian Museum of Natural History, Budapest; IVB, Institute of Vertebtrate Biology, Brno; NMK, National Museums of Kenya, Nairobi; NMP, National Museum Prague, Prague; ROM, Royal Ontario Museum, Toronto; TCWC, Biodiversity Research and Teaching Collections (formerly the Texas Cooperative Wildlife Collection), College Station; TM, Ditsong National Museum of Natural History (formerly Transvaal Museum), Pretoria; UADBA, Université d’Antananarivo, Départment de Biologie Animale, Antananarivo; ZFMK, Zoological Research Museum Alexander Koenig, Bonn. Table S2. Eigenvector loadings of the principal components analysis (PCA) for principal component (PC) 1 and PC2 based on standardized craniodental measurements of the species traditionally placed in Neoromicia sensu Simmons (2005) and in Laephotis s.s. Table S3. Eigenvector loadings of the principal components analysis (PCA) for principal component (PC) 1 and PC2 based on standardized craniodental measurements of the Neoromicia capensis group (see main text for the species included). Table S4. Uncorrected Cytb p-distances among (below diagonal) and within (numbers on diagonal) 53 Vespertilionidae species calculated in MEGA X v.10.0.5. Figure S1. Maximum likelihood phylogeny of mitochondrial cytochrome b sequences of 418 sequences of Vespertilionidae specimens. Bootstrap values are adjacent to nodes. Specimen localities include counties for Kenya. DRC refers to Democratic Republic of the Congo, and CAR refers to Central African Republic. Museum acronyms are defined in the Material and Methods section. Sequences downloaded from GenBank are indicated by inclusion of GenBank accession numbers (Supporting Information, Table S1). Figure S2. The type locality of Pseudoromicia kityoi on the edge of Mabira Forest (photograph by S. Waswa Babyesiza).

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